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Full text of "The smaller fossil Foraminifera of the Panama canal zone"

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SMITHSONIAN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 
Bulletin 103 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



ON SOME FOSSIL AND 

RECENT LITHOTHAMNIEAE OF THI. 

PANAMA CANAL ZONE 



By MARSHALL A. HOWE 
Of the New York Botanical Garden 



Extract (rom Bulletin 103, pages 1-13, with Plato 1-11 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

I9I8 



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ON SOME FOSSIL AND RECENT LITHOTHAMNIEAE OF 
THE PANAMA CANAL ZONE. 



By Marshall A. Howe, 

Of The New York Botanical Garden. 



INTRODUCTION. 

The following report is based chiefly upon a number of specimens 
of fossil calcareous algae, of the group known to geologists as " Nulli- 
pores," from Oligocene and Pleistocene strata in the Panama Canal 
2^ne, collected in 1911 by D. F. MacDonald and T. W. Vaughan, of 
the United States Geological Survey. 

In this material the Pleistocene period is represented by a single 
collection (MacDonald, 6039), consisting of numerous excellent free 
specimens, " from flats near Mount Hope, five feet above tide level.'' 
These Pleistocene specimens appear to the writer to belong to a 
species found by him a year or two earlier to be living in the Colon 
region, only a few kilometers distant. This species, so far as the 
writer can determine, has been hitherto undescribed ; in framing its 
diagnosis, as published below, the fossil as well as the recent mate- 
rial has been considered, but a recent specimen, being more complete 
and satisfactory for detailed study, has been named as the technical 
type of the species. 

So far as the present writer has been able to discover, the fossil 
coralline algae of America, in their taxonomic aspects at least, offer 
a practically untouched field for research. It is, of course, possible 
that geological and paleontological papers in which calcareous algae 
have been described have escaped the attention of phycologists, but 
inquiry among American geologists and paleontologists and a search 
of accessible literature have thus far revealed to the writer but a 
single * hitherto described species of fossil Lithothamnieae from the 
Western Hemisphere, namely, Lithoth/mmium curasavicum K. Mar- 
tin, from the Island of Curasao, a species to which further allusion is 
made below in the discussion of ArehaeolithotJiamnium episporum. 

^ Stromatopora compacta Billings (Palaeozoic Fossils, vol. 1, p. 65, 1862) from the 
Island of Montreal, etc., has sometimes been considered by geologists to be of coralllna- 
ceons affinities (the species has been referred to Solenopora by Nicholson and Btherldge, 
GeoL Mmg^ vol. 8, p. 529, 1885), but, if we may judge from published figures, the organ- 
ism seems to the writer hardly a coralline alga, if indeed it is an alga at all. 



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2 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

The fossil Lithothamnieae of Europe have been described and fig- 
ured in considerable number and with various degrees of care and 
detail. Most of these European descriptions and figures the writer 
has been able to see; some of them offer a reasonable basis for the 
future recognition of the forms concerned, without a reexamination 
of the original materials, but many of them do not. The present 
writer has had access to a good representation of the living Litho- 
thamnieae of North America, the West Indies, Europe, and the East 
Indies, but so far as the fossil forms are concerned, he has had to 
depend upon descriptions and figures alone, which, as stated 
above, are often very unsatisfactory. In venturing to propose as 
new, two species of Lithothamnieae from Oligocene strata of the 
Panama Canal Zone, he doubtless risks the possibility that some 
future investigator, working with better materials or even with the 
same, may be able to convince himself or even to prove conclusively, 
that one or both of said species should be considered identical with 
species previously described from Europe. The diagnostic charac- 
ters, the limits of variation, and the geographic range of even the 
living species are still very imperfectly understood. Some of the 
species are evidently widely distributed within certain temperature 
limits; others are at present known from single localities. So far as 
may be inferred from our present knowledge, very few, if any, of the 
forms of Lithothamnieae now living in tropical America occur also 
in European waters. 

List of Species and Theib Geologic Occurrence. 

ArchaeoUthothamnium episporum, new species. Recent, Toro Point; and 
Pleistocene, Mount Hope; both in the Canal Zone. 

LiihotJvamnium vaughanii, new species, Oligocene, Cnlebra formation at 
station 6026, about half way between Monte Llrio and Bohio Ridge. 

Lithothamniwn isthmi, new species, Oligocene, Emperador limeBtone at 
stations 6021, about 4 miles north of Gamboa Bridge, and 6024-6, Rio 
Agua Salud, Panama Railroad (relocated line). 

Lithoporella melohesioides (Foslie) Foslie, Oligocene, Emperador lime- 
stone at station 6024-c, Rio Agua Salud, Panama Railroad (relocated 
line). 

ARCHABOUTHOTHAlfNnJM ^ EPISPORUM, new ipedes. 

Plates 1 to 6. 

Brownish red when living, the thallus forming at first widely ex- 
panded crusts 0.25-1.0 mm. thick, these in many cases repeatedly 
overgrown, the resulting crusts becoming 6 mm. or more thidc, some- 
times remaining nearly smooth or exhibiting the irregularities of the 

^ We follow Rothplets'8 original spelllDg of the final syllable of this unfortunately long 
name, a spelling that, happily, agreer with Philippics spelling of the final syllable of 
Lithothamnium, 



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GEOLOGY AND FALEOKTOLOGY OF THE CANAL ZONE. 8 

substratum alone, but more often developing coarse, irregular 
rounded excrescences 5-12 mm. in diameter, or short rounded ver- 
rucae or nodules 2-5 mm. in diameter, the surface in sterile parts 
mostly smooth, indurated, and occasionally subnitent; hjrpothallia 
varying from weakly to strongly developed, 30-170 |x thick, their 
cells 17-28 pi by 8-11 ii; cells of the perithallium in distii^ct and regu- 
lar layers except in oldest and youngest parts, the layers in more or 
less distinct zones, layers of short and of long cells occasionally al- 
ternating, cells mostly 8-15 \l by 5-8 jjl, in decalcified condition sub- 
monUiate, sphaeroidal to ellipsoidal, 1-2| times as high as broad, 
in calcified condition mostly subquadrate or oblong in vertical sec- 
tion: sporangia superficial, their apicula even with the surface, or 
slightly protruding, their cavities becoming only imperfectly and 
insularly embedded, the sori slightly elevated, very irregular in 
outline, mostly 0.1-1.0 mm. broad, often widely confluent and anas- 
tomosing and becoming 5 mm. or more broad, the surface at length 
whitish and scarious, the ostioles mostly 16-22 pi in diameter, sporan- 
gia 65-96 |i high (including apiculum), 27-50 pi broad, 4-partite (oc- 
casionally 2-partite?), the spores irregularly paired or rarely sub- 
zonate. 

Localities and geologic occwrrence. — Covering dead corals, etc., 
and often forming concretionary pebbles with coral cores, from low^ 
water mark to a depth of several meters. Point Toro, near CJolon, 
Panama Canal Zone, Howe 6882 (type, in Herb. N. Y. Bot. Gard.), 
January 7, 1910 ; Colon, Howe 6840 (this covers continuously a mass 
of old coral 32 cm. long and 14 cm. in greatest width) ; also, as a 
Pleistocene fossil, " from flats near Mount Hope, five feet above tide 
level,'' D. F. MacDonald, station 6039,^ 1911. 

Paratypes.—CHt. No. 35298, U.S.N.M. 

In outward form and in its habit of overgrowing old corals, 
Archaeolithoth^mwmtm episporum resembles A. erythraeum (Roth- 
pletz) Foslie, f. durum (Heydrich) Foslie, from the Red Sea and the 
East Indies, especially as illustrated by Weber- van Bosse and Foslie 
(Corallinaceae of the Siboga Expedition, pi. 5). Of this species we 
have seen only one specimen (from near Makassar), communicated 
by Mme. Weber- van Bosse, but from this and from the descriptions 
Mid figures of A. erythraeum published by Foslie, Heydrich, and 
Lemoine, we infer that the Panamanian specimens represent a differ- 
ent species. Perhaps the most important distinctive character of A. 
episporum, is to be found in its more superficial sporangia, as may be 
seen by comparing our photographs (pi. 2, fig. 1; pi. 3) with Hey- 
drich's figure * of a vertical section through a sporangial sorus of his 

^This is associated with minor amoanta of other crostaceous corallines, including 
tAthophylWm, species, and OonioUthon, species. 
» Ber. Dents. Bot Ges.. toI. 15, p. 68. flg. 2. 1897. 



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4 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Sporolithon ptychoides^ which Foslie ^ and Lemoine ^ consider to be 
synonymous with A. erythraeum. The sori or the emptied sporangia! 
cavities appear also to be much less regularly embedded or overgro\^n 
by new tissue than is the case in A. erythraeum^ if one may judge 
from Rothpletz's original description,* Heydrich's figure 3,* Le- 
moine's figure 2&,* and the descriptions given by the last-named writ- 
ers; however, Foslie*^ remarks of ^1. er^ythraeum that "the sori are 
partly to be found overgrown in great numbers by new formed tissue, 
partly, however, they are not to be seen in section." In A. episporum^ 
the sporangia themselves have never been seen except close to the 
surface; the emptied sporangial cavities do not show in a rough frac- 
ture or in an ordinary ground section, but irregular traces of them 
are often to be found in thin microtome sections of decalcified ma- 
terial. The sori of A. episporv/m are so superficial that their cover- 
ing, after the discharge of the spores, appears to die and is flaked off 
together with more or less of the intersporangial parts, and the new 
tissue growing up from the base of the sorus shows only occasionally 
and imperfectly the outline of the former sporangial cavities. 

Kothpletz's original description of his LUhothamnium erythraevum 
leaves one in some doubt as to whether he found the contents of the 
sporangium divided or undivided; he uses the term " Tetrasporen," 
but the measurements that he gives for these " Tetrasporen " are 
such as commonly belong to the whole sporangium in this group. 
In Heydrich's first description * of his Sporolithon ptychoides^ the 
" Tetrasporangien " are said to be "meist ungetheilt, selten zwei- 
theilig," but a little later " he figures four tetraspores in a sporangium, 
arranged in the " cruciate " manner. But this mode of division being 
at variance with the prevailing ideas as to the arrangement of the 
spores in the Corallinaceie, Foslie,** a little later in writing a diag- 
nosis of the genus ArchaeoUtJiotharrmiimi inserted a question mark 
after "sporangia * * * unparted or cruciate?" and this sign of 
doubt as to the cruciate division has been repeated by later writers.* 
In A. episponmi the mature sporangia are commonly and normally 
4-parted in an irregularly " cruciate " fashion, but often the division 
axes of the two pairs of spores are at right angles to each other, so 
that only three spores are visible in a lateral view, and occasionally 



^ Sihoga Exped. Monog., No. 61, p. 38. 1904. 

* Ann. Inst Oc^nog., yoL 2, pt 2, p. 67. 1911. 

* Rottapletz, A. Bot Centralb., vol. 64, p. 5. 1898. 
♦Ber. Dents. Bot Ges., vol. 15, p. 68. 1897. 
*Sihoga Exped. Monog., No. 61, p. 41. 1904. 
•Ber. Dents. Bot Ges., vol. 15, p. 69. 1897. 
'Idem, pi. 18, fig. 3. 

•Kgl. Norske Vldensk. Selsk. Skr. 1900, pt 5, p. 8. 1900. 

•De Toni, Syll. Alg., vol. 4, p. 1721, 1905; Svedelius, in Bng. k Prantl, Nat. PflanKen- 
fam., vol. 1, pt 2; Nachtsttge, p. 267, 1911. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 5 

the second divisions seem to be omitted and the sporangium is appar- 
ently mature with only two spores. Vexry irregular types of division 
also occur, and rarely one finds an approach to the zonate ^ arrange- 
ment characteristic of most of the Corallinaceae. 

The perithallic cells of A. episporv/m appear to be, in the decal- 
cified state, more rounded and in more moniliform filaments than is 
^e case in A. erythoHtewm^ as may be seen by comparing our photo- 
micrograph* with the photomicograph of a presumably decalcified 
section of A. erytkraeum — ^published by Lemoine.' The distinct strat- 
ification of the perithalliiun of A. epi^porwm is due, in part, to the 
alternation of layers of long and short cells, but we have never seen 
in the Panamanian species any such striking alternation of long and 
short cells as is shown in this photograph published by Mme. Le- 
moine and as is shown still more emphatically in Heydrich's figure 3 ^ 
of a vertical section of his Sporolithon ptychoides. 

From ArchaeoUthothamninm dimotum Foslie and Howe,* the only 
living species of this genus previously described from the West 
Indian region, A. epispormn differs widely in its thicker crusts, in 
its more superficial sporangial sori, which are for the most part ex- 
foliated after maturity of the sporangia and are only obscurely and 
imperfectly overgrown, in the usually larger, more rounded, and 
more moniliately arranged cells of the perithallium, the larger and 
rather less widely separated sporangial ostioles. etc. 

ArchaeolUhotJiamnium curasavieum (K. Martin) Foslie,* a Creta- 
ceous fossil from the island of Curasao, is described and figured as 
showing distinctly rows of embedded sporangial cavities, such as 
would not be seen even in a thin decalcified section of A. episporum. 

A Pleistocene fossil, collected by MacDonald at station 6039, from 
flats near Mount Hope, came from a few kilometers from the locali- 
ties where we found the plant living, and we can entertain no serious 
doubt as to the specific identity of the recent and the fossil forms. 
The living and fossil are similar in external habit^ as may be seen 
by comparing plates 1 and 4. They are similar also in their rela- 
tions to old corals, and in structure (compare fig. 1, pi. 2, and fig. 4, 
pi. 5) they appear to exhibit only such differences as may be ascribed 
to individual variation or as may be expected in comparing the recent 
or living with the long dead. But little remains of the fossil speci- 

>Z<mately 4-parted sporangia have been described by Foslie for the Callfomlan 
ArehaeoUthoihainntmm tonatotporum (Foslie, Algologiske Notiser. II. Kgl. Norske 
Vidensk. Selsk. Skr., 1906, pt 2, p. 14), so that It would appear that this larenus exhibits 
A wide Tarietj in the matter of division of its sporangia. 

« Plate 3, fig. 2. 

•Ann. Inst. Oc^noj?., vol. 2, pt. 2, pi. 1, flg. 1. 1911. 

« Ber. Dents. Bot. Oes., vol. 15. 1897. 

• BnIL N. Y. Bot. Gard., vol. 4, p. 128, pi. 80, flg. 1 ; pt. 87, 1906. 

^ lA$hothamnium euratavicum K. Martin, Berictat liber eine Reise nach Niederl&ndisch 
WesMndlen nod daranf gegrfindete Stndlen. II. Geologle, p. 26, pi. 2, flgs. 22-25, 1888. 



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6 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

mens after decalcification, though the outlines of the cells may be 
recognized here and there. As microtome sections of the decalcifie<i 
fossil material are out of the question, comparisons of structure of 
the recent and fossil must naturally be based upon calcareous groimd. 
sections. And in comparing the cell structure in sections of the re- 
cent decalcified specimens (pi. 3) with that shown in ground sections 
of the calcareous fossils, it is necessary, of course, to bear in mind 
that cells in calcareous ground sections of the Corallinaceae com- 
monly appear much more rectangular than in decalcified sections af 
the same material.^ In the sections of the fossil material thus far 
made there are no certainly recognizable traces of sporangial cavi- 
ties, but this is true in almost an equal degree of calcareous 
ground sections of the recent specimens except as to the surface of 
the plant (fig. 1, pi. 2), where the sori are, in fact, so decidedly 
superficial or even exserted that they could, perhaps, hardly be 
expected to persist in the fossil state. 

In the same locality with the type-specimens (Howe 6832) there 
occurs an outwardly somewhat Similar plant (Howe 6837) that we at 
first suspected to be the antheridial form of A, episporum^ but certain 
recognizable, though possibly unimportant, differences in the form, 
size, and zonation of the perithallic cells have restrained us from so 
considering it. The antheridial conceptacle (cavities) in this 6837 
are 64-95 ^ broad and 60-72 ji high ; they become copiously embedded 
by the continued upward or outward growth of the thallus. 

UTHOTHAMNIUM > VAUGHANII, new ipeclei. 

Plate 7, figs. 1 and 2, and plate 8. 

Thallus forming at first expanded crusts 1-2 mm. thick, these be- 
coming overgrown, irregularly stratified, and 10 mm. or more thick, 
developing finally nmnerous, rather coarse, crowded anastomosing 
branches, and forming masses 2-4 cm. or more high ; branches mostly 
3-12 mm. in diameter, usually much flattened, occasionally subterete, 
often reduced to anastomosing ridges, or sometimes appearing as 
dome-shaped elevations 2 cm. or more broad; primary hypothallia 
somewhat reduced, their cells 14-33 ^ by 8-14 pt, rather irregularly 
arranged (i. e., not distinctly "coaxial"), cells of medullary hypo- 
thallia mostly 15-30 ^ by 5-13 pi, secondary hypothallia numerous 
and thin ; branches showing in section numerous narrow irregularly 
flexuous, often subelliptic-lenticular or subcrescentic zones caused by 

1 For illustrations of these differences, see Lemoine, Ann. Inst Oc^nog., vol. 2, pt. 2, 
p. 45, flgs. 19-21, 1911. 

2 The writer believes, with Mme. Paul Lemoine, that the current rules of nomenclature 
require that Philippics original spelling of this generic name should be respected, eren 
though prevailing usage has modified the final syllable. Whether the rules of nomencla- 
ture Justify the use of this generic name for any of the species now bearing it is a more 
complicated question. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 7 

the alternation of layers of short and long perithallic cells, or by the 
interpolation of reduced secondary hypothallia ; the larger perithallic 
cells mostly 18-62 \l by 11-14 |x, usually higher than broad, the 
smaller subquadrate, about 8 (i. square, or sometimes much compressed 
(7 |& high, 14 \L broad) ; conceptades becoming embedded; tetrasporic 
conceptacles much flattened, oblong or elliptic-oblong in radio-ver- 
tical section, the cavity 500-740 |x in maximum width, 130-230 \l in 
height ; roof of the tetrasporic conceptacle rather sharply defined, its 
cells in regular vertical rows of 1-4 cells, often elongate vertically, 
becoming sometimes 25-80 (jl high. 

Loedlity and geologic occurrence. — Oligocene, Culebra formation, 
^about half way between Monte Ldrio and Bohio Bidge, on the relo- 
cated line of the Panama Kailroad," collected by D. F. MacDonald 
and T. W. Vaughan, 1911 (station No. 6026). 

Holotype and paratypes.—Cst. Nos. 35299, 35800, U.S.N.M. 

The specimens obtained are more or less embedded in a hard rock 
matrix, so that our photograph (fig. 1, pi. 7) can give only an imper- 
fect idea of the outward form of the plant. With a little mental 
clearing away of the matrix, it seems probable that in size and ex- 
ternal appearance, the species may be compared with rather coarse 
eroded conditions of the living Lithothanvniv/m glaciale Kjellman, 
but there is little similarity in structure; the perithallic cells of Z. 
vaughanii average considerably larger than those of L. glaciale and 
they are arranged in more distinct layers; the embedded tetrasporic 
conceptacles of L, vaugham^i are more flattened than those of Z. 
glaciale^ their cavities have about twice the maximmn width of those 
of L. glaciale and the specialized character of the conceptacle roof 
is not noticeable in Z. glaciale. 

In external habit Lithotha/nmium vaughanii may perhaps be com- 
pared also with the living LithopkyUum racemua (Lamarck) Foslie 
forma crassvm (Philippi) Foslie^ of the Mediterranean and Ad- 
riatic seas, especially as shown in Hauck's figure 2 * under the name 
Lithotlummiwm, crasaum Philippi, though the Panamanian fossil 
sometimes develops longer and perhaps more flattened branches than 
this form. 

Of the living Lithothamnieae now known to the present writer as 
occurring in the West Indian region, Lithothamninm vaughanii per- 

^KgL Norske VldenBk. Selsk. Skr., 1898, pt. 8, p. 9, 1898. Foslle*8 Identlflcatioii of 
lAthothomniMm onutum Philippi as a form of lAthophyllum raoemus (Lamarck) Foslie 
was accepted by Heydrlch (Bot Jahrb., vol. 23, p. 586, 1901), bat Mme. Lemolne qnofes 
JAthoihamnium oraa$um Philippi as a synonym of lAthothamnium caloareum (Pallas) 
Areschong. It is not, however, apparent that any of these writers examined authentic 
BKaterlal of Phillppi's Lithoihamnium ora89um, if such exists. It is of some interest, also, 
to note that less than six months before Heydrich accepted LttTiothamnium orassum 
PhUlppi as a form of lAthophyllum racemuM he named it as the type of a proposed new 
gtams SUehotpora (Ber. Dents. Bot Gtes., vol. 18, p. 816, 1900). 

* Haack, P. Die Meeresalgen Deutschlands und Oesterreichs. In Rabenhorst, L.» 
Kryptogamen-Plora von Deatschland, Oesterreich und der Schweiz, vol. 2, pi. 1, 1885. 



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8 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

haps most i*esembles Lithophyllum daedaleum Foslie and Howe* as 
to general habit, but differs from it much in structure. 

In the best section, No. 35299 U.S.N.M., the one from which the 
photographs (fig. 2, pi. 7 and pi. 8) were made, the coarse inter- 
sporangial sterile tissue of the tetrasporic conceptacles is scarcely 
shown, yet the roofs of the conceptacles show unmistakable canals 
and none of the conceptacles in section exhibits a single orifice, so 
that we consider ourselves justified in inferring that the specimen in 
(luestion is tetrasporic and that it belongs in the genus LithotJuimr- 
nium in the sense in which that name is currently applied to living 
plants. In a section from another specimen under the same collec- 
tion number, traces of the sporangia and of the intersporangial 
sterile tissue are evident. It is to be observed also that the zonate 
arrangement of tissues, as observed in a section, is essentially of the 
character assumed by Mme. Lemoine ^ as being peculiar to the genus 
Lithothamnium. The rather distinctly specialized nature of the 
cells of the conceptacle roof is evidently a character of importance, 
in which respect it differs markedly from the plant we are de- 
scribing as LitJwthamniuvi isthmi^ as also in the distinctly zonate 
structure of the thallus, the reduced hypothallium. the larger tetra- 
sporic conceptacles, larger perithallic cells, etc. 

Among the more fully described fossil Lithothamnieae, L, vaagh- 
anil may perhaps be compared with Lithothamnium suganum Roth- 
pletz^ from the Tertiary (" Scio-Schichten ") of Val Sugana, near 
Borgo in the Austrian Tyrol, but the conceptacles of the Panamanian 
fossil are much larger (500-740 jjl wide and 130-230 |jl high vs. 250 [k 
wide and 100 jjl high) and the perithallic cells appear to average con- 
siderably larger, being sometimes 13-22 \l high, while those of L. 
suganum are described as 9-12 |i. long. 

UTHOTHAMNIUM ISTHBQ. new ipMie*. 

Plate 7, fig. 3; plates 9. 10, and 11. 

Thallus forming at first stratified crusts 3-12 mm. thick, but at 
length developing tortuous anastomosing branches and forming large 
rather solid, concrescent, fruticose masses; branches mostly 2-12 mm. 
in diameter, much flattened or subterete, often subconic-cylindric^ 
tlexed-digitiform, or molariform; hypothallia showing regular con- 
centric layers of cells ("coaxial") ; hypothallium of the crustaceous 
parts 160-480 \i thick, its cells 17-28 p. by 8-13 pi, transition to the 

1 Bull. N. Y. Bot. Gard., vol. 4, p. 133, pis. 83, 84, 93, 1906. 

* Lemoine, Mme. Paul. Structure anatomlque des M^lobesl^es. Application k la classl- 
0cation. Ann. Inst. Ocdanog., vol. 2, pt. 2, pp. 27, 28, 1911. 
»ZeIt8. Deuts. Geol. Ges., vol. 43, p. 319, pi. 17, fig. 4, 1891. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 9 

perithallium abrupt ; medullary hypothallium of the branches mostly 
0.6-2.0 mm. in diameter, often turning yellow and more or less disin- 
tegrated, its cells 17-44 jjl by 8-18 (i, transition to the perithallium 
abrupt or gradual; cells of the perithallium in distinct layers, the 
layers in rather indistinct zones; perithallic cells of the crustaceous 
parts subquadrate, 8-11 \l in diameter, sometimes only 6 \l broad; 
perithallic cells of the branches usually a little higher than broad, 
8-19 |i by 8-12 \l ; conceptacles becoming embedded ; tetrasporic con- 
ceptacles appearing much flattened in a vertical section, the cavity 
240-550 \L in maximum width, 130-105 \k in height. 

Localities and geologic occurrence. — In Emperador limestone of 
Oligocene age (and often constituting the dominant element in its 
composition) on relocated line of the Panama Eailroad, opposite San 
Pablo, Panama Canal Zone (" first limestone outcrop just north of 
Caimito Station, about four miles north of Gamboa Bridge"), col- 
lected by D. F. MacDonald and T. W. Vaughan, 1911, Station No. 
6021 (No. 35301, type) ; and " above foraminiferous marl at Agua 
Salud Bridge about ^ mile north of New Frijoles on relocated line, 
Panama Eailroad," by the same collectors. Station No. 6024i. 

Holotype and paratypes.—Csit Nos. 35301 to 35303, U.S.N.M. 

The material upon which the above description is based shows 
much variation in form and structure and it was our first impression 
that two or more species were represented in it. However, if this is 
true, the two or more species are so intergrown and entangled and 
are so similar in structure that it is difficult to determine where one 
begins and the other ends. As regards the vegetative structure, we 
believe that we have been able to trace the continuous organic con- 
nection of the two types shown in our photomicrographs (pi. 9 and 
fig. 2, pi. 11), yet it is notoriously easy in the case of overgrowing 
and overgrown fossil Lithothamnieae to mistake the close contact of 
independent plants for structural continuity. 

In the tetrasporangial specimen (No. 35301 — fig. 3, pi. 7 and pi. 9) 
that we have named as the type, the thallus presents itself in the 
form of irregularly superposed crusts, more or less overlaid by 
crusts showing a somewhat different structure and conceptacles of 
a different sort, these outer layers probably representing a crustace- 
ous species of Lithophyllum. The hypothallium of this No. 35301 is 
suggestive of that figured by Foslie^ for his living TMhothamnmm 
fragilissimum from Borneo (which, however, has a much thinner 
thallus) . It suggests also the hypothallium of LitJiothamnium lich- 

^Foslie, M. Lithothamnioneae, Melobesieae, Mastophoreae. In Weber-van Bosse, A., 
and Foslle, M. Tbe CoraUlnaceae of the Siboga Expedition, Siboga Exped. Monog. No. 61, 
fig. 6, 1004. 



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10 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

enoides^ as figured by Rosanoff * and by Lemoine,' but the crusts 
evidently more massive than in that species. 

Although the outward form of Lithothamnium isthmi is more 
less obscured by being embedded in rock, it seems probable that in 
typical condition (No. 35301) the external appearance of the ph 
may be compared with the recent plant from the Adriatic Sea fi^i 
by Hauck^ as '^ LitJiophyllum decussatum Solms,'' which Fosl 
afterwards referred to his LithotJumvnium phdUppii — a species tl 
he maintained even after conceding* its specific identity with i 
earlier-published LithophyUvm crispatum Hauck. The typi* 
form of Foslie's Lithothamniurrh phUippu is said by him • " to ha 
its hypothallium distinctly marked and vigorously developed, for 
ing a coaxilate layer," but the "coaxial" character is essentia! 
denied by Mme. Lemoine^ to what she considers the same spec 
under the name LUhothammmri crispatum Hauck. The perithal 
cells of the crustaceous parts of Lithothamnium isthmi appear 
average considerably smaller than those of L, crlspatuvi {L. pi 
lippii) according to the measurements given by Lemoine and 
Foslie, The tetrasporangial conceptacles of the LithophyUum a 
cussatum of Hauck (Lithothamnium phiUppii Foslie) are stated 1 
Hauck to be " 800|iL bis 1 mm.'* in diameter, while in Lithothamniu 
isthmi they arc only 240-550 p. in maximum width. Moreov< 
unless we are mistaken in connecting the fruticulose parts of t 
Panamanian fossil with the crusts, Lithothammium isthmi develo 
numerous solid anastomosing branches, while in Z. crispaMtm tl 
short branchlike excrescences are mostly hollow, infundibulifori 
or scyphiform. These fruticulose conditions, which comprise 
large part of the material collected by MacDonald and Vaugha 
suggest in external form certain stat^ of the living West Indii 
LithophyUum daedaleum Foslie and Howe, which also presen 
itself in both crustaceous and fruticulose conditions. Occasional 
an unusually long subterete branch may resemble in form a f ra 

^M4in. Soc. Imp. Scl. Nat. Cherboarg, vol. 12, pi. 6, flg. 14, 1866. 

* Ann. Inst. Oc^anog., vol. 2, pt 2, flg. 60, 1911. It Is of interest to note that Mn 
Lemoine, basing her system of classification primarUy npon the Tegetative stmctnre 
the thallus, leaves LitJiothamnium UohenoMes in the genus lAthoph^llum, notwithstai 
Ing the fact that its tetrasporangia are borne as in the genus LUhothamMum of mode 
writers. In the same way she would doubtless place lAthottMtmnium Uthmi \n the gen 
LithophyUum, even though this species (or its type at least) clearly has the tetrasport 
glal conceptacles of the conventional Lithothamnium. 

'Hauck, F. Die Meeresalgen Deutschlands und Oesterreichs, pi. 1, flg. 7. See al 
pi. 1, flg. 1, of FosUe's Die Lithothamnien des Adriatischen Meeres und Marokkos (Wit 
Meeresuntersuch, Helgoland, vol. 7, pt 1, 1904). 

« Foslie, M. On some Lithothamnia. Kgl. Norske Vidensk. Selsk. Skr., 1897, pt. 
p. 7, 1897. 

* Wiss. Meeresuntersuch, Helgoland, vol. 7, pt. 1, pp. 18, 14, 1904. 
•Kgl. Norske Vidensk. Selsk. Skr., 1900, pt. 1, p. 6, 1900. 

* Ann. Inst Octenog., vol. 2, pt 2, p. 80, flg. 88, 1911. 



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qboijOot and palboktolooy of the oakal zone. 11 

m^it of the living East Indian Lithothamnmm ptUchrum A. Weber 
and Foslie.* 

LitJiothammufri fosUei (Trabucco) De Toni (Syll. Alg., vol. 4, p. 
1761, 1905), a Miocene fossil from Italy, is figured* as having a 
** coaxial ^ hypothallium, but from the illustrations given of the con- 
eeptades, there is no sufBcient ground for considering this plant 
to be a LU7u>t?uimnium rather than a Lithophylhmi. In the original 
place of publication nothing but a figure (section) is given, from 
which, according to the scale of magnification given, it would appear 
that the conceptacles are only 140-160 (x by 80-90 (& and the perithallic 
cells about 16 (x high, making the cells rather larger and the con- 
ceptacles much smaller than in L. isthim. 

If we are correct in including with Lithothammum isthrm the more 
ramified forms collected by MacDonald and Vaughan, the species, 
thou^ commonly coarser, appears to be sometimes suggestive of 
plants figured as NulUpora rajnosissima Reuss or LUhothmmdvm 
ramosissimum (Reuss) Schimper, from the Tertiary " Leithdkalk " 
of the vicinity of Vienna, but Eeuss's original figures and descrip- 
tion* relate to external form only, and give no adequate basis for 
referring the plant to a modem genus. Unger^ adds good figures 
of the vegetative structure, but shows no conceptacles. Rothpletz " 
describes the conceptacles of L, ramosissimum as 280 (& high, while 
the height of the conceptacles of L. isthmi is 130-165 (i and the width 
240-550 v^ Rothpletz has no doubt that there are two species of 
Lathothamnieae in the '^ Leithakalk," which may have been confused. 

UTHOPOBELLA MELOBESOmES (FosUe) Fo«ll«. 
Lithoporella melohesioidcs (Foslle) Foslie, Kgl. Norske Videusk. Selsk., 

1909, pt. 2, p. 50. 
Mastophora melobesioides Fosluc, Kgl. Norskd Vldensk. Selsk. Aarsber., 

1902. p. 24, 1903. 

Locality and geologic occurrence, — This species occurs in small 
quantity with LUhotJummium iatJmd in Emperador limestone of the 
Oligocene age, "above foraminiferous marl at Agua Salud Bridge 
abont i mile north of New Frijoles on relocated line, Panama Rail- 
road," D. F. MacDonald and T. W. Vaughan, 1911, Station No. 60246. 

EXPLANATION OF PLATES. 

Plate 1. 

Archaeolithothamnium episporum M. A. Howe. 

Photograph, natural size, of the type-speclinens, collected at Point Toro, near 

Colon, Panama Canal Zone. January 10. 1910 (Howe 6832). The technical 



1 Compare pi. 4. aibodra Bzped. MoDog. 61. 

* BuUthothamniHm Jo$HM, Trab. BoU. Soc Qeol. Ital., vol. 19, pi. 11, fig. 10, 1900. 

* HaWHngtr, Katurw. Abh., t<H. 2, pt. 1, p. 29, pi. 8, flgi. 10, 11, 1848. Type from 
* Naodiyrfl,'* Hungary. 

* Denksehr. k. Akad. Wiss. Wien, toI. 14, p. 23, pi. B, flgs. 18-22, 1868. 
"Zeits. Dents. OeoL Oes., vol. 48, p. 820, 1891. 

8870*— ISh—Bull. 108 2 



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type in a narrower sense is tlie specimen shown at die lower riglit-liand comer 
of the plate — the specimen from which figure 1 of plate 5 was obtained. 

Plate 2. 

ArcJiaeolithothamnium episporum M. A. Howe. 

Photographs of radio- vertical ground (calciferous) sections of type mateirlal 
(Point Toro, Howe 6882). 

J^^G. 1. Lithothamnium vaughanii M. A. Howe. Photograph of the type- 
portion of a sporangial sorus, enlarged 42 diameters. 
2. Section, enlarged 200 diameters. 

Plate 3. 

Archaeolithothamnium epispoi^m M. A. Howe. 

Photographs of radio-vertical sections of decalcified material (Point Toro, 
Howe 6882), enlarged 200 diameters. 

Fig. 1. Section showing sporangia and tetraspores. 

2. Section showing emptied sporangia, form and arrangement of peri- 
thallic cells, a weakly developed hypothallinm, etc. 

Plate 4. 

Archaeolitfiotfiainnium epispomm M. A. Howe. 

A Pleistocene fossil, " from flats near Mount Hope, five feet ahove tide level,** 
D. F. MacDonald 6039, 1911, natural size. 

Plate 5. 
Arohaeolithothamnium epiapoi^m M. A. Howe. 

Figs. 1 and 2. Photographs of the type material (Point Toro, Howe 6832). 

Fig. 1. Portion of the surface, showing the more or less confluent sporangial 
sori, enlarged 4 diameters. 
2. A smaller part'^of the same surface, showing the sporangial ostioles, 
etc., enlarged 25 diameters. 
Figs. 3 and 4. Photographs of Pleistocene specimen from Mount Hope (Mac- 
Donald, Cat No. 35298, U.S.N.M.) 
Fig. 3. Radiovertical section showing several superposed crusts and three well- 
developed hypothallia, enlarged 42 diameters. 
4. A part of a cross section of one of the excrescences or hranches, showing 
a single weakly developed hypothallinm, enlargetl 42 diameters. Oom- 
pare structure of living specimen as shown in fig. 1, plate 2. 

Plate 6. 

Archaeolithothamnium epispomm M. A. Howe. 

Photograph of section of the Pleistocene fossil from near Mount Hope 
(Cat. No. 35298. U.S.N.M.). A section magnified 78 diameters. 



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OBOLOGT AND PALEONTOLOGY OF THE OANAL ZONE. 13 

Plate 7. 

Fia. 1. Lithothamnium raughanii M. A. Howe. Photograph of the type- 
Bpecimens (between Monte LIrio and Bohio Ridge, MacDonald and 
Vaughan, Cat. No. 352d9, U.S.N.M.), natural size. 

2. Lithothamnium vaughanii, A section showing irregular zonatlon, tetra- 

sporic conceptacles, etc., enlarged 42 diameters. 

3. Lithothamnium isthmi M. A. Howe. A section, slightly enlarged (11/8 

of the natural dimensions), showing the type-specimen embedded in 
the nintrix (from about 4 miles north of Gamboa Bridge, MacDonald 
and Vaughan, station 0021). The type material (CJat. No. 36301. 
F.S.N.M.), from which the section shown in plate 9 was obtainetl. 
occupies the centnil portion of the light area and is overgrown by 
crusts of what appears to be n different plant, probably a species of 
LithophyUnm. * 

Plate 8. 

Litti^tha milium vaughanii M. A. Howe. 

An enlargement of a part of the section shown in figure 2, plate 7, illus- 
trating form of peri tha 111c cells, the reduced secondary hypothallia, the some- 
what specialized roof of the tetrasporic conceptacles. etc. Magnification 300 
diameters. 

Plate 9. 

Lithothamnium isthmi M. A. Howe. 

A section of the type material (MacDonald and Vaughan, station 6021, Oat. 
No. 86301, U.S.N.M.), enlarged 100 diameters. The section shows the well- 
deTeloped "coaxial" hypothalllum, the smaller-celled perithallium, and the 
conceptacles with the coarse interspornngial tissue characteristic of the genus 
Lithothamnium, 

Plate 10. 

Lithothamnium isthmi M. A. Howe. 

A specimen from about one-third mile north of New Frljoles (MacDonald 
and Vaughan, station 6024-&, Cat. No. 85305, U.S.N.M.), natural size, showing 
fofltil embedded in matrix, In both weathered and freshly broken surfacef). 

Plate 11. 

Lithothamnium isthmi M. A. Howe. 

A somewhat obliquely transverse section of a branch (specimen from about 
4 mfles north of Gamboa Bridge, MacDonald and Vaughan, station 6021. Oat. 
No. 8S802, n.S.N,M.), enlarged 106 diameters. 



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BULLETIN 103 PL. I 




ARCHAEOLITHOTHAMNIUM EPI8P0RUM M. A. HOWE. 
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BULLETIN 103 PL. 2 





Archaeolithothamnium episporum M. a. Howe. 

For explanation of platk sii paoi |2. 



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BULLETIN 103 PL. 3 




^::m'^m3^i^j^^};C9j- :■■■ 




Archaeolithothamnium episporum M. a. Howe. 

For explanation of plate see page 12. 



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BULLETIN 103 PL. 4 




Archaeolithothamnium episporum M. a. Howe. 

For explanation of platb tee pace 12. 



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BULLETIN 103 PL. 6 





^jft^.^-. 







2 4 

Archaeolithothamnium episporum M. a. Howe. 

FOft eXPUANATION OF PLATE BEE PAOE 1 2. 



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BULLETIN 103 PL. 6 




Archaeolithothamnium episporum M. a. Howe. 

For explanation of plate see page I2. 



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BULLETIN 103 PL. 7 






I. 2. LlTHOTHAMNIUM VAUQHANII M. A. HOWE. 3. LiTHOTHAMNIUM ISTHMI 

M. A. HOWE. 



Fon EXPLANATION OF PLATE SEE PAGE 13. 



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BULLETIN 103 PL. 8 




LITHOTHAMNIUM VAUQHANIi M. A. HOWE. 
For kxplanation of plate see paqe 13. 



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BULLETIN 103 PL. 9 




LITHOTHAMNIUM ISTHUI M. A. HOWE. 
For explanation of plate see paqe 18. 



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BULLETIN 103 PL. 10 




LITHOTHAMNIUM I8THMI M. A. HOWE. 
For explanation op plate see PAoe 13. 



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LITHOTHAMNIUM I8THMI M. A. HOWE. 

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INDEX. 



Page. 

ArcbaeoUtbottaamnliim 4 

dlmotum — — 5 

episporom — 1-6, 12 

erythraeum 3-5 

zonatosporum 6 

EiilithotluLmnlum fosllel 11 

Gonlolltlion 8 

ritbophyllum 3. 9-ill, 18 

crlspatQin lOj 

daedaleum 8, 10 

decussatum 10 

racemus 7 

Idtboporella melobeaioldee — . 2,11 

Liltbotlminnlum 2, 8, 10, 11, 18 

craBStun 7 

curasaylcnm 1, 5 



Page. 

Lithothamnium fosliei 11 

fragilissimum U 

gladale 7 

lathmi 2, 8, 10, 11. 13 

lichenoides 9, 10 

phiUppU 10 

pnlchnun 11 

ramoissimaiii 11 

■uganum 8 

vaoghanU 2, 6-8, 13 

liastopbora melobesioides . 11 

Nulllpora ramoBissima 11 

Sporolithon ptychoides 4, 5 

Stichospora 7 

Stromatopora compacta 1 



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SMITHSONIAN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 
Bulletin 103 



Contributions to the geology and f 

totogy of the canal zone, panama 

geologically related areas in c 

tral america and the west indil 



» 



THE FOSSIL HIGHER PLANTS FJ 

THE CANAL ZONE 



By EDWARD W. BERRY 
Of the Johns Hopkins University, Baltimore 



Estract from Bulletin 10% pages 15-44, with Plates 12-18 






WASHINGTON 

COVERNMENT PRINTING OFFICE 
1918 



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SMITHSONIAN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 
Bulletin 103 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



THE FOSSIL HIGHER PLANTS FROM 
THE CANAL ZONE 



By EDWARD W. BERRY 
Of the Johns Hopkint University, Baltimore 



Extract from BuUetio 103, page$ 15-44, with Platc$ 12-18 




WASHINGTON 

GOVERNMENT PRINTINO OFHCE 

1918 



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THE FOSSIL HIGHER PLANTS FROM THE CANAL ZONE. 



By Edwabd W. Berry, 
O/ the John% Sopkint University, Baltimore. 



INTRODUCTION. 

It is a truism that the present floras and faunas of Central America 
are the result of a long series of antecedent geologic changes which 
mig^t be amplified as geographic, climatic, and biologic. As the 
past can only be understood by means of our knowledge of the 
present, so, too, the present can only be understood by means of 
our knowledge of the past. Moreover, this can never be a local prob- 
lem, and this is particularly true of the Isthmus of Panama marking 
as it does at times the highway of communication between the ter- 
restrial life, both animal and plant, of North and South America; 
at other times marking one of the paths of communication between 
the marine life of the Atlantic and Pacific. Thus the history of the 
Central American region is of the utmost importance in any con- 
sideration of the extinct terrestrial faunas and floras of North Amer- 
ica or the marine faunas that formerly flourished on the east and 
west coasts. 

Our knowledge of the present flora of the isthmian region is based 
upon Seemann's flora * and Hemsley 'S flora of Central America, sup- 
plemented by the scattered papers by numerous authors on special 
topics relating to this flora. As the results of the recent Biological 
Survey of the Canal Zone become available, we will doubtless have 
a secure basis for comparisons with antecedant floras both in this 
regi<m and the areas north and south of it. 

The present distribution of plant associations is in its broader 
outlines governed almost entirely by the interrelations between 

1 R. T. Hill, who did some geological work on the Isthmos in 1806 for Alexander 
^g*y**- mentions lignite and fragments of fossil plants in the Cnlebra clays at the base 
of t& canal catting at Cnlebra station (Bull. Mas. Comp. Zool., yoi. 28, No. 5, 1808), and 
the Ilgnltlc coal at Chiriqai Lagoon was studied by Dr. John Evans in 1857, who reported 
** that the fossil plants associated with the coal were endogenous and allied to or identical 
with those at present growing in the ylclnity." (Repts. of Expl. & Sury. for the Loca- 
tion of Inter-oceanic ship canals, etc., by the U. S. Naval Bxped., 1876, B. P. Lull, 
V. & N.,' commanding, Washington, 1879.) 

'Seemann, Flora Panamensis. Botany of the voyage of H. M. S. Herald, pp. 57-846, 
3862-1867. 

15 



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16 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

topography and the prevailing winds and the resulting variations 
in rainfall. 

The climate is now moist tropical, modified by the nearness of 
the two oceans, and there is therefore but slight diurnal or JtmnifrT 
variations in temperature. So far as information is available re- 
garding the conditions during the Tertiary, there is no evidence that 
can be deduced from the fossil flora or the geographical history of 
the region to indicate that the climate was very different from what 
it is now at any time during the Tertiary, unless we are prepared to 
assent to enormous changes in the altitude of the land, for which 
the data does not seem to be adequate. 

The prevailing winds now come from the northeast, and as the 
divide is near the Pacific Coast the major part of the Isthmus north 
of this low divide has a heavy rainfall, as, for instance, 170 inches 
at Porto Bello and 129 at Colon, as compared with 90 inches at 
Culebra or 71 inches at Ancon. There are two seasons — a short 
relatively dry season extending from January to April and a long 
and relatively wet season the balance of the year with the maximum 
of precipitation from September to December. Before the clearing 
of the French Canal Company forests covered six-tenths of the 
Isthmus, the remainder being broken forests and savannas. Ever- 
green tropical rain- forests of mixed angiosperms covered the entire 
northern watershed and part of the Darien region on the south side. 
Some of the forests of the southern watershed are what are known 
as monsoon forests, with many deciduous species, and at high alti- 
tudes there may be more gregarious types of forest as, for example. 
the oak forests which are so striking a feature in the uplands of 
Central America as you proceed to the northwest. 

The shores are skirted with dunes abounding in Leguminosae and 
Euphorbiaceae with. Coco palms and Hippomane. Low shores and 
tidal inlets are covered with mangrove swamps with Rhizophora, 
Avicennia, Conocarpus, etc. Less saline coastal marshes are covered 
with Acrostichum, Crescentia, or Paritium thickets. The evergreen 
forest is composed chiefly of species of Sterculiaceae, Tiliaceae, and 
Mimosaceae, Euphorbiaceae, Anacardiaceae, Bubiaceae, Myrtaceae, 
and Melastomataceae, with small palms like Chamaedorea, Trithri- 
nax, and Bactris. 

CORRELATION. 

The fossil flora described in the present report is too limited*for 
purposes of exact correlation, which may be expected to be settled 
by the marine f aimas present at most horizons in the Isthmian region. 
Begarding the plants in the various f ormational imits recognized in 
the Canal Zone by MacDonald a glance at the accompanying table 
of distribution will show that from the oldest (Bohio) to the young- 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 17 

^st (Gratun) plant-bearing formations there is no observable dif- 
ference in floral fades, and while the plants are entirely too few for 
positive conclusions, and while not much variation can be expected 
in fossil floras of the Tropics unless after the lapse of long intervals 
of time or the intervention of marked changes in physical conditions, 
T am disposed to think that this so-called Oligocene series of forma- 
tions does not represent any great interval of time. 

Nearly all of the fossil plants are new, the only outside occurrences 
being the Hieronyrfda which is common to the Tertiary of Ecuador 
and the -Po&TWWjy&m and Taenioxylon both of which occur in the 
Oligocene of the island of Antigua, and both have related types in 
the Oligocene (Catahoula and Yicksburg) of our Southern States 
In addition to the Hieronyrma common to Ecuador there are several 
other elements in the Tertiary flora of the latter region that are 
similar to Panama forms, and it is not improbable that the coals of 
Loja in the Ecuadorian Andes are the same age as the so-called 
Oligocene series of Panama. Only one pre-Oligocene plant is re- 
corded from Panama and the age (Eocene) rests on the stratigraphic 
observations of Doctor MacDonald and paleontologic determinations 
by C. W. Cooke. The form itself offers no intrinsic evidence of its 
age and might well be early Oligocene but for the fact that Doctor 
MacDonald collected the type stratigraphically below a bed con- 
taining a varietal form of the moUusk, Venericardia planicosta, 

Th^ chief question of interest in the correlation of these Panama 
beds is their equivalence in terms of the European section. The 
present flora offers no evidence on this point which must hence be 
tletermined by the accompanying marine faunas. However, in view 
of the traditional unscientific assumption that all of the f ossilif erous 
beds of the Carribbean region are Oligocene in age, it is of interest 
to note that Douvill6* from a study of the foraminifera, pointed 
out as early as 1898, that a considerable part of the so-called Oligo- 
cene of the Isthmus was Aquitanian and Burdigalian in age; that is 
to say, lower Miocene according to the present conceptions of Euro- 
pean geologists and palentologists. 

In my preliminary announcement * of the discovery of fossil plants 
in the Canal Zone I stated that none of the plants recognized indi- 
cated Eocene and that they were all probably Oligocene in age. 
This statement was perhaps overemphasized in a desire to offset the 
extreme views of certain foreign paleontologists who have held that 
these faunas were young Miocene or even Pliocene. 

The question of the exact time in the Tertiary at which connections 
between North and South America were i-eplaced by marine condi- 
tions is of the utmost importance in all studies of distribution of both 

^ DoutUI^, H. BhU. Soc Q€ol, de France, ser. 8, yoL 26, pp. 587-600» 1898. 
* Bwrj, B. W. Science, new ser., yol. 89, p. 857, 1914. 



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18 



BULLETIN 103. UNITED STATES NATIONAL MUSEUM. 



the marine faunas and the terrestrial faunas and iSoras. The floral 
evidence as previously stated is inconclusive. I should not, however, 
be inclined to consider any of the fossil plants, except one Eocene 
species, described in the present report as younger than Burdigalian 
nor older than Sannoisian (Lattorfian). 



P^lmotglon palmacUts. 



J^ieui euMnenait. 



OuatUrta euUhremi* 

MpriitoeophpUum panamtme. . 



TaenhMjflon mulHradiatum . 
Inffa oliffoeaeniea 



Cditia euM»rerui9 

Airoea oligoeaeniea 

BanitUrta praenuntia. 
Hierofiifmia Uhmanni. 



tekmidelia beJuuntU 

Metpttodavkne euUhrentit : 
Cilfptrantha gatunentit , . . 
MeliitomUes mieonioida. . , 



Diotvvrot maedonaldi. 
MondeUtia ooldmani. . . 
BubiadU* izoreoidet, . . 



6840 
X 



6840 
X 



6840 
X 



6846 
X 



6623 
X 



6845 
X 



6837 
X 



6837 
X 



6837 
X 



e I 



6S86b 
X 



Pftlmrays * X J mile 8. of Empirt 

6887 Bridge. 

Femfragmento 6837 (/. Aeroitlcbaiii. 

BOTANICAL CHARACTBR. 

The fossil flora at present known from the Canal Zone is extremely 
limited and entirely too small for either purposes of adequate cor- 
relation or for deductions concerning the true botanical f acies or the 
environmental conditions. Seventeen species are determined and 
two or three additional forms are tentatively recognized. This 
paucity is especially to be regretted since it is improbable that under 
the existing climatic conditions as favorable opportunities for the 
discovery and collection of fossil plants will ever be presented as 
during tiie digging of the canal. While fossil plants were nowhere 
found to be abundant in the shales, nevertheless, it is very probable 
that an experienced collector by working over a large amount of 



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GBOLOOY AND PALEONTOLOGY OF THE CANAL ZONE. 19 

material could have gotten together a much more representative col- 
lection. 

The plants collected include ill-defined fragments of one fern, 
two nndertermined species of palm, represented by fragments of 
foliage, and a third represented by petrified stems, and 16 dicotyle- 
dons, of which two are represented by fruits and the balance by 
leaves. 

Among the Dicotyledonae there are representatives of the orders 
Urticales^ Ranales^ Resales^ OeranidUSj SapindaleSy ThymelealeSj 
Myrtales^ EbenaleSj and Rubiales. Orders conspicuous in the exist- 
ing flora of the Isthmian region unrepresented among the fossils are 
the AraleSy Poalesj CyperaleSy and OrchiddUs among the Monocoty- 
ledonae, and the Campanulales and Peraonales among the Dicoty- 
ledonae. 

The following 14 families are represented by fossils in Panama: 
Moraceae, Anonaceae, Myristicaceae, Mimosaceae, Caesalpiniaceae, 
Papilionaceae, Malpighiaceae, Euphorbiaceae, Sapindaceae, Laura- 
ceae, Myrtaceae, Melastomataceae, Ebenaceae, and Bubiaceae. Only 
the last, with two species, is represented by more than a single species. 
When so sparse and evenly distributed a representation of the 
families is present in a fossil flora, it is an indication that after allow- 
ing for some accidents of preservation, those families represented may 
be regarded as the most abundantly represented in the Tertiary flora 
of the region, and in this respect there is a very great similarity to the 
existing flora of the Isthmian region. The present forests of Panama 
are made up principally of species of Arecaceae, Moraceae, Mimo- 
saceae, Papilionaceae, Sterculiaceae, Tiliaceae, Euphorbiaceae, Ana- 
cardiaceae, Myrtaceae, Melastomataceae, and Bubiaceae. The only 
ones of this list not found fossil are the Sterculiaceae, Tiliaceae, and 
Anacardiaceae, and as these three families are all abundant in the 
much more complete floras from the Tertiary of the southeastern 
United States, it is safe to assimie that they were also present in the 
Tertiary flora of Panama. The mainly herbaceous families abundant 
in the Recent flora, which are hardly to be expected in the fossil flora, 
are the Poaceae, Qrperaceae, Orchidaceae, Araceae, and Compositae. 

The bowers of wild figs of the existing flora are represented by a 
small-leafed species of Ficus from two localities in the Culebra 
formation. The family Anonaceae, which has numerous species of 
Anona and Gtiatteria in the Becent flora of Central America, is rep- 
resented by a fine large species of the latter genus which is not un- 
common in the Gatun, Caimito, and Culebra formations. Guatteria 
contains about 50 existing species of tropical shrubs and trees of 
varying habitats and exclusively American, and has not been previ- 
ously recognized with certainty in fossil floras. Anona is abundant 



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20 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

in the Eocene and Oligocene of our Southern States, but Ouatteria 
has not been recognized. 

The Myristicaceae is represented by an infrequent species of 
Myriaticophyllwm in the Culebra formation, and in this connection it 
is of interest to note the presence of fruits and seeds of Myristica in 
the uppermost Eocene of Texas suggestive of the subgenera Virola 
and Compsoneuray both of which occur in the Recent flora of Central 
America. The Leguminosae have three fossil species. The Mimosa- 
ceae, which are very abundant in the existing forests of Panama, are 
represented by a fossil species of Inga^ a large genus of tropical trees 
with upward of two-score species in Central America, nearly half 
of which are recorded from Panama. Inga is well represented in 
the abundant Eocene floras of our Southern States, and it is of in- 
terest to note the resemblance between the fossil species from Panama 
and a species described by Engelhardt from an unknown Tertiary 
horizon in Ecuador. 

The Caesalpiniaceae is represented by a single species of Cassia^ a 
large genus not only in the Recent equatorial floras but well repre- 
sented in most fossil floras from the Upper Cretaceous to the present. 

The Papilionaceae, very abundant in the existing flora of Panama, 
is supposed to be represented by the petrified wood of a large tree 
referred to the genus Taenioxylon and found in the Cucuracha, 
Culebra, and Bohio formations. 

The family of Malpighiaceae is represented by the genera Hiraea 
and Baniateria. The former has about 30 recent species, exclusively 
American, ranging from Mexico and the Antilles to tropical Brazil 
and Peru, and it is represented by a fossil species in the Eocene of 
the Mississippi embayment Banisteria contains about 80 existing 
species, mostly climbing shrubs. It is at present confined to the 
American tropics, but appears to have been present in Europe as well 
as in the southern United States during the Tertiary. 

The Euphorbiaceae, abundantly represented in the present forests 
of Panama, is represented in the Caimito formation by a species of 
Hieronymia apparently identical with one described by Engelhardt 
from the Tertiary of Ecuador. Hieronymia^ not otherwise known 
in the fossil state, contains about a dozen existing species which are 
confined to tropical America, where they range from Mexico and the 
West Indies to Brazil. 

The Sapindaceae, abundant in all fossil floras from the Upper 
Cretaceous onward, and exceedingly abundant in the Tertiary floras 
of the Mississippi embayment, is represented in the fossil flora of 
Panama by a species of Schmidelia found in the Caimito and Culebra 
formations. Sch/nddelia has a large number of existing species in 
the equatorial regions of both hemispheres and, except for petrified 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 21 

material from the island of Antigua, it has not previously been recog- 
nized in the fossil state. 

The family Lauraceae, so extensively represented in the Tertiary 
floras of the Mississippi embayment and in the Becent tropical flora 
of South America, is represented at Panama by a single fragmentary 
species which is referred to Mespilodaphne. The latter has numer- 
ous modem species in the tropics of America and Africa. 

The Myrtaceae, one of the abundant families in the existing for- 
ests of tropical America, has a fossil species of CaJyptranthes at 
Panama. This genus has about 70 exclusively American existing 
species ranging from Mexico and the West Indies to southern Brazil. 
Hemsley records 7 recent species from Central America, of which 
2 are foimd on the Isthmus. It is also represented in the lower 
Ek>cene of the Mississippi embayment. The abundant, both Becent 
and fossil, representatives of the allied genera Eugenia and Myrda 
have not been recognized in the fossil flora of the Isthmus. 

The Melastomataceae, an immense tropical family in the existing 
flora and very abundant throughout Central America, has a single 
fossil species in the Culebra formation. 

The family Ebenaceae, usually abundant in fossil floras from the 
Upper Cretaceous onward, and with a large number of species in 
tropical Ajnerica, is represented on the Isthmus by the petrified 
fruits of a species of ebony {Diospyros) known to be from an older 
horizon (Eocene) than the balance of the known fossil flora. 

The Rubiaceae, a prominent family in the existing flora of Central 
America, where according to Wallace (1911) it ranks fourth in size 
with 146 species, is represented by two fossil species, both found in 
the Gatun formation. These are referred to Rondeletia and Ruhi- 
acUes, 

The former has not heretofore been found fossil. It includes 
about 70 existing species of a variety of habitats, confined to the 
American tropics and chiefly massed in the Antilles and Central 
America. RvhicBcites is represented by a fruit which is apparently 
referable to the tribe Ixoreae, now confined to the tropics of both 
hemispheres. 

TERTIARY ECOLOGY. 

The restricted variety and fragmentary condition of the fossil 
plants thus far collected inhibits a detailed discussion of the prob- 
•able ecology of the Tertiary flora. In so far as climatic conditions 
are concerned the Tertiary plants indicate an abundant rainfall and 
relatively high equable temperatures such as prevail at the present 
time in the Hill country and Coastal Plain of the Isthmus. There is 
no indication of upland vegetation. None of the fossil plants indicate 



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V 
22 BULLBTIK 103, UNITED STATES NATIONAL MUSEUM. 

mountains sufficiently high to harbor that mixture of temperate types 
such as is seen at the present time in the mountains of Central 
America, as, for example, above 6,000 feet in Costa Rica. There was 
plenty of opportunity for the introduction of such types had the 
climate been propitious, so that I would infer that the Tertiary relief 
was slight, that is under 5,000 feet and probably much less than this, 
although there is no evidence to warrant precision of statement. 

On the other hand, the collected floras do not furnish any traces 
of the characteristic vegetations of low muddy shores, although types 
like Bhizophardj Avicenma^ Conocarpus^ Lagimcuiaria^ etc, were 
already in existence in Eocene times as we know from their presence 
in the Mississippi embayment of that time, where they were undoubt- 
edly derived from the south. I do not infer that these costal types 
were absent in the Tertiary flora of the Isthmus. On the contrary 
they must have been present; but no traces of them have been dis- 
covered except the traces of Acroatichum in the Culebra formation. 

The bulk of the fossil plants clearly belong to the evergreen rain 
forests and they have the appearance of having been washed into 
the basins of sedimentation by streams. None of the lithologic speci- 
mens that I have seen from the Isthmus indicate autochonous swamp 
deposits either of coastal or valley situations and I picture the flora 
as one of a humid tropical character covering a country of low hills. 
This is of necessity a tentative conclusion and perhaps even such 
general deductions are unwarranted because of the very limited data 
with which I have had to deal. 

FLORA OF THE CANAL ZONE. 

Arecales: 

Arecaceae — 

Pahnoxylon palmacites (Sprengel) Stenzel. 
Urticales: 

Moraceae — 

Ficus culebrenaisj new species. 
Ranales : 

Anonaceae — 

Guatteria culebrenais^ new species. 
Myristicaceae — 

Myristicophyllum panamense^ new species. 
Resales : 

Leguminosae — 

Taenioxylon multiradiatufn Felix. 
Inga oUgocaemcoj new species. 
Cassia culebrenris^ new species. 



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QEOLOOY AND PALEONTOLOGY OF THE CANAL ZONE. 2H 

Geraniales: 

Malpighiaceae — | 

HircBa oligocaenica^ new species. 
Banisteria praenuntia^ new species. 
Euphorbiaceae — 

Hieronyrnda lehmanni Engelhardt? 
Sapindales : 

Sapindaceae — 

Schmidelia hejucenMs^ new species. 
Thymeleales: 
Lauraceae — 

MespilodaipJme cuiebrensis^ new species. 
Myrtales: 

Myrtaceae — 

Calyptranthes gattmensts^ new species. 
Melastomataceae — 

MelastomiteB micomoides^ ney^ species. 
Ebenales : 

Ebenaceae — 

Dioapyroa maodondldif new species. 
Rnbiales: 

Rnbiaceae — 

Rondeletia goldmam^ new species. 
Rubiacitea ixoreoidesj new species. 
Fern fragments of Acrostichum. 
Palm rays. 

SYSTEMATIC PALEOBOTANY. 

PTERIDOPHYTA. 

Order FILICALES. 

FERN FRAGMENTS OF ACROSTICHUM. 

The material from the Culebra formation, one-fourth mile south 
of Empire Bridge, contains several obscure fragments of large simple 
fern pinnules with reticulate venation strongly suggestive of Acros- 
tichum, but too incomplete for identification. The genus now prin- 
cipally represented by the cosmopolitan tropical tidal marsh species 
AcrosUchv/n^ auremn, is abundant in the Eocene and Oligocene of 
both America and Europe, and is especially characteristic in the 
Jackson, Catahoula, and Vicksburg of our Gulf States. 



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24 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

SPERMATOPHYTA. 

Order ARECALES. 

Family ARECACEAE. 

PALM RATS. 

The broken rays of apparently two species of palms occur spar- 
ingly in the Cul^ra formation at the locality one- fourth mile south 
of Empire Bridge. These are too incomplete for even tentative 
generic determination. 

Genus PALMOXTLON Schenk. 

Group LUNARIA. 

PALMOXTLON PALMACrTBS (Sprang^l) Stonad. 

Plate 12. fig. 1. 
EndogefUtes palmacites Spsenoel, Ck>mmentatio, p. 39, flgsu 6, 6a, 1S28. 
FaacicuUtes palmacites Cotta, Dendrol., pp. 49, 89, pi. 9, figs. 1, 2, 1832. — 

Unoeb in Martius, p. 59, tab. geol. 3, fig. 6, 1845. 
Palmacites dubius Ck>BDA, Beitrage, p. 42, pi. 22, 1845. — Schikpeb, PaL 

V6g^t., vol. 2. p. 513. 1870; Handbuch, Abst. 2, p. 887. 1892. 
Palmncylon tenerum Felix, Foss. H51zer Westindiens, p. 26, pi. 4, fig. 1, 

1883.— Schenk in Zlttel. 
PtUmoxyUm palmacites Stengel, Foss. PalmenhOlzer, p. 245, pi. 20, fig. 25S, 

1904. 

Description. — Fibro- vascular bundles small, very numerous, closely 
spaced, orbicular or ovate in cross section, imifonnily distributed as 
a rule, 0.60 mm. to 0.75 mm. in diameter, and rarely, if ever, that 
distance from oua another. Auxiliary bundles absent 

Sclerenchyma portion excavated more or less deeply to receive the 
vascular portion, which is often nearly equal to it in size. Occa- 
sionally a thin zone of sclerenchyma entirely surrounds the vascular 
portion. Sclerenchyma fibres small, isodiametric, greatly thickened, 
of nearly uniform size, about 0.035 mm. in diameter. Vessels vari- 
able in size, ranging from 0.072 mm. to 0.18 mm. in diameter, usually 
two large vessels and either none or several small vessels on the side 
away from the bast in each bundle. The phloem portion in general 
destroyed and represented by a disorganized cavity between the ves- 
sels and the bast. 

The ground mass of the st^n consists of thin walled parenchyma 
without intercellular spaces. The cells are small, isodiametric, 
rounded pentagonal or hexagonal except where there are but one or 
two rows between closely adjacent bundles, in which case they are 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 25 

narrowly compre^ed and elongated parallel to the sides of the 
bundles. Their diameter varies from 0.035 mm. to 0.10 mm. Scat- 
tered through the stem parenchyma are darker cells which in polar- 
ized light appear to be gum cells. They are slightly larger than the 
parenchyma cells, being from 0.072 mm. to 0.108 mm. in diameter. 

Occasional bundles are seen to be branching. These are the fas- 
ciculi fhroduGtores or Kreuzungsbiindel. 

This species was first recognized by Sprengel in 1828, who referred 
it to Endogenites; Cotta four years later transferred it to Fasciculites, 
and Corda in 1845 referred it to Pdlmacites. When Felix came to 
publish on the Antigua woods in 188S he recognized this species, 
but in describing it under the genus Pahnomylon which had been 
proposed by Schenk only a year or ^wo before he took the liberty 
of giving it the new name of tenerumj which under the rules of 
nomenclature has no standing as Stenzel recognized in print in 1904. 

The specimen from Panama is small and may be from near the 
periphery of a stem, although in the group Lunaria there is little 
diflPerence between the central and peripheral regions. In the size, 
outline, and crowding of the fibrovascular bundles as well as in the 
character of the parenchyma of the groundmass the present species 
greatly resembles Pdhnoxylon integrum described by Felix ^ from 
Cuba and considered by Stenzel ^ as merely a variety of the Antiguan 
species Pcdmoxylon antiguense (Unger) Felix.' It differs from that 
species in altogether lacking the numerous auxiliary sclerenchyma 
bundles which are so well marked in Paknoxylon integrum. A fur- 
ther difference is the presence of gum or mucilage cells which are 
fairly numerous in the Panama specimen of Paimoxylon palmaoites 
and which might upon a merely superficial examination be mistaken 
for auxiliary sclerenchyma bundlea Among the Oligocene species 
of Pdhaoxylon from the southern United States Palmoxylon miaais- 
iippense Stenzel* is very similjir to the present species. 

Other described fossil species which show more or less resem- 
blances are Palmowylo-n stellatum^ aschersoni^ variabiles and ceylani- 
cum. The nearest aflinity among recent palms is not determinable 
in the present state of our knowledge of the anatomy of the latter. 
The present type of structure is commonly known as the C!ocos- 
like type. 

For some unknown reason the upper Eocene and lower Oligocene 
in southeastern North America abounds in silicified palm wood. 
Palm leaves are often very abundant in the Wilcox and Claiborne 
Eocene and in the Apalachicola Oligocene; but all of the petrified 

1 Felix, Fobs. HOla. WestliidlenB, p. 24, pi. 5, flg. 2, 1883. 
> Stensel, Fobs. PalmenhOlser, p. 154, pi. 1, figs. 1-10, 1904. 
• Felix, Fobs. HOls. Westlndiens, p. 22, pi. 4, flg. 5. 
« Steniel, Fobs. PalmenhOlser, p. 248, pi. 21, flgs. 264-265. 



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26 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

palm wood in our Gulf region is confined to the Jackson or Vicks- 
burg groups. 

The island of Antigua, celebrated for at least a century for its 
petrified woods, has furnished at least seven species of petrified 
palms, five of which were known to Unger as eariy as 1850, and one 
was figured by Witham in 1833. These also are of Oligocene age. 
There are two additional Oligocene species described from the West 
Indies without definite information as to exact locality, and there 
is also a species from Trinidad and another from Cuba. The Oligo- 
cene species at present known from the southern United States are 
seven in number, four of which have not been found outside of that 
region, while one or possibly two are common to Antigua, and a third 
has been reported by Felix from Southern Mexico. 

Occurrence. — Cucuracha formation, green clays, Gaillard Cut (loc 
6586). Collected by D. F. MacDonald. 

Collection.— TJ, S. National Museum. Cat. No. 35310. 

Order URTICALES. 

Family MORACEAE. 
Genus FICUS Unnaeus. 

FICUS CULEBBEN8IS, ii«w speclM. 

Plate 13, tig, 1. 

Description. — ^Leaves of relatively small size, broadly oblong- 
lanceolate in general outline, apex acute but not extended or cuspi- 
date. Base bluntly pointed. Margins evenly rounded. Texture 
coriaceous. Length about 8 cm. Maximum width, in the middle 
part of the leaf, about 2.15 cm. Petiole short, stout, and curved. 
Midrib stout and prominent on the under surface of the leaf. Sec- 
ondaries thin, very numerous, evenly spaced, subparallel; they di- 
verge from the midrib at wide angles averaging about 75 degrees, 
pursue an almost straight outward course, their ends being con- 
liccted well within the margins by regular flat arches formed by 
their abrupt camptodrome endings. Tertiaries obsolete. 

This is an especially well-marked species of the lanceolate leafed 
section of Ficua^ and it may be matched by a number of still exist- 
ing species found in the American tropics. Among such a large 
number of both existing and fossil forms detailed comparisons are 
not especially pertinent. Two comparisons that seem significant are 
the resemblance of the present form to Ficus newtonensis Berry of the 
Upper Claiborne of the Mississippi embayment and to the forms from 
the Sannoisian of Haering in the Tyrol which Ettingshausen ^ refers 

^ EttiDgshausen* Tert. Fl. von Haering, p. 41, pi. 10, figs. 6, 8, 1858. 



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OBOLOOY AND PALEONTOLOGY OF THE CANAL ZONE. 27 

to Fieus jynx Unger, but which appear to me to be decidedly diflfer- 
ent from Unger's type. 

Occurrence, — Culebra formation, upper part. East wall of the 
GaiUard cut just north of Canal Commission station 1760 (collected 
by M. I. Groldman). 

Order RANALES. 

Family ANONACEAE. 
Genus 6UATTERIA Ruiz and Pavon. 

GUATTESIA CULEBHENSIS. new ipMies. 

Plate 13, fig. 2. 

Description, — ^Leaves of large size, broadly ovate in general out- 
line, with a narrowed slightly decurrent base and a narrowed and 
extended acuminate tip. Length about 20 cm. Maximum width, 
approximately midway between the apex and the base, between 6 cm. 
and 7 cm. Margins entire. Texture coriaceous. Petiole short and 
stout, enlarged proximad, about 2.25 cm. in length. Midrib stout 
and promient. Secondaries mediumly stout and prominent, about 
ten opposite to alternate pairs diverge from the midrib at angles 
ranging from 45*^ to 60°, sweeping upward in regular ascending 
subparallel curves, camptodrome in the marginal region. Tertiaries, 
where visible, percurrent. 

The present is one of the more abundant and better preserved 
forms from the Canal 2iOne, but the large size of the leaves usually 
results in fragmentary specimens, the tip being almost invariably 
missing. The species shows great similarity with various existing 
forms of Anonaceae. It is very close to Anana majrcgravii Martins of 
Venezuela, French and Dutch Guiana, and Brazil (Bahia and Per- 
nambuco). It is, however, among the various species of Guatteria 
that the closest homologies are found. The latter genus contains 
about fifty species of shrubs and trees, exclusively American^ and 
found in Mexico, Central America, tropical South America, and in 
the northern Andes. The fossil may be compared with a large num- 
ber of the existing species, as for example GhiaUeria ouregon Dunal, 
a large tree of the Carribbean islands and equatorial South America, 
Guatteria dolichopoda De CandoUe or G. grandiftora De CandoUe 
of Central America. 

The family Anonaceae contains about 700 existmg species, dis- 
tributed among about 48 genera, only two of which are present in 
North America. The family is practically confined to the Tropics, 

>The Asiatic species of yarious authors are referred to the genus Polyalthia, 
8370*— 18g— Bull. 103 3 



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28 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

a single Australian species, and the North American g^us Aaimiiia^ 
with 6 or 7 species being the only conspicuously extratropical forms. 
The area of maximum representation is southeastern Asia and the ad- 
joining region of Malaysia, for while only 16 genera are confined to 
this region it contains over 350 species, and six additional genera 
{Miliusa^ Vvaria^ PolyaltJda^ Oxymitra^ Melodorwm^ and Poporvia) , 
with a total of over 250 species have the bulk of their species in this 
area. Only a single genus is confined to Australia, and the bulk of 
the Australian species are to be regarded as migrants from the pre- 
ceding area. There are upwards of 100 species and 6 peculiar genera 
in tropical Africa; and America has about 200 species and 10 peculiar 
genera. These are all confined to the Tropics, except for a species of 
Anona^ which reaches the coast of peninsular Florida, and for the 
genus Asimina^ with six or seven species of shrubs and small trees of 
the south Atlantic and Gulf States. One of these, Asimina triloba 
Dunal, is hardy as far north as New York, and has the distinction 
of growing the farthest distance from the Equator of any existing 
member of the family. The fossil record of the Anonaceae is very 
incomplete, only the genera Anona Linneaus and Asimina Adanson, 
being known with certainty. Both of these genera are present in 
the flora of the Wilcox group of the Mississippi embayment. 

The genus Chiatteria has not, so far as I know, been heretofore 
found fossil, except for a doubtful species described by Hollick from 
the Upper Gretaceous of Marthas Vineyard and Long Island. The 
genus Uvaria Linnaeus has a Pliocene and three Pleistocene species 
on the Island of Java, and the genera Melodomim Dunal and Mitre- 
phora Blume are both represented in the Pleistocene of that island. 

The genus Anona has from fifteen to twenty fossil species, five of 
which are also represented by seeds. The oldest is a species described 
from the Dakota sandstone. There is a second species in the late 
Cretaceous or Early Eocene of the Rocky Mountain province. The 
flora of the Wilcox affords a glimpse into the true stage of evolution 
of Tertiary floras in that expanded belt of the American equatorial 
region which was the center of radiation of so many recent types. 
There were three exceedingly well-marked species of Anona alon^ 
the Wilcox coast and their leaves are very common at some localities, 
although no seeds have as yet been discovered. I assume that these 
Wilcox forms had habits similar to those of the majority of the ex- 
isting species, exemplified by our Florida Anona glabra Linnaeus, 
or pond apple, which frequents shallow fresh-water swamps, low 
shady hammocks, or stream borders near the coast. Other species 
occur in the low coppice association or on edges of brackish swamps 
on the Bahamas. The cultivated species, as, for example, the Ameri 
can Anona reticiilata Linnaeus, which is planted in Guam, often 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 29 

spreads naturally along the inner beaches, while attempts to intro- 
duce others of the most highly esteemed American species in the 
Orient have failed. From its prevalence among the existing species 
the habit of growing in wet, shaded soils is evidently an old one, and 
since the Wilcox Anonas are associated with a strand flora the as- 
sumption that they grew on the inner beaches or the shaded and 
more swampy edges of lagoons possesses every degree of probability. 

In the pipe days of Alum Bay which were contemporaneous with 
the Wilcox there are two species of Anona^ and Engelhardt has de- 
scribed two species from the Eocene or Oligocene of Chili. The 
Oligocene record shows a species in France and a second in Saxony. 
In the Miocene there are two species each in England, Styria, and 
Croatia, and one each in Bohemia, Colorado, and Transylvania. 
There is one each in the Pliocene of France and Italy, showing how 
modem was their extinction in the south of Europe. 

The genus Asimina has only four or five recorded fossil species. 
These are all American except for a form from the Pliocene of Italy 
which has been referred to this genus, although I suspect that it 
represents Anona^ since Asimina appears to have originated and been 
confined to the Western Hemisphere. The oldest known species is 
based on foliage which is found in the basal Eocene of the Eocky 
Mountains (Denver formation) and of the embayment (Midway 
Group). There is a single species based on a seed from the basal 
Wilcox and no other records except a fono close to the modem 
from the late Miocene of New Jei'sej'^ (Bridgeton sandstone) and the 
occurrence of the existing Asindna triloba Dunal in the interglacial 
beds of the Don valley in Ontario. There are 17 existing species of 
Anona recorded from Central America, six of which are known from 
Panama. Hemsley records 11 species of Guatteriu from Central 
America, at least two of which occur in Panama. 

Occurrence. — Culebra formation, upper part. East wall of Gail- 
lard Cut just north of Canal Commission station 1760 (collected by 
M. L Gk)ldman). Gatim formation. Gatun borrow pits (collected 
by M. I. Goldman). 7 miles northeast of Bejuca near Chame 
(=Caimito formation) (collected by MacDonald). 

Family MYRISTICACEAE. 
Genus MYRISTICOPHYLLUM Geyler. 

MTSISTICOPHYLLUM PANAMEN8E, ii«w spcdM. 

Plate 18. fig. 3. 

Description. — Leaves ovate or ovate lanceolate in outline with 
pointed apex and base, entire, evenly rounded margins, subcoriaceous 
in texture. Length about 9 cm. Maximum width, midway between 
the apex and the base, about 3.3 cm. Petiole slender, about 8 mm. 
long. Midrib slender. Secondaries thin, about 8 subopposite ascend- 



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30 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

ing subparallel pairs; they diverge from the midrib at acute angles 
and are subparallel with the lower lateral margins, eventually canip- 
todrome. Tertiaries obsolete. 

This species is unfortunately represented by fragmentary remains 
inadequate for conclusive identification. The genus Myristica Lin- 
naeus contains about two score existing species, rather more than half 
being American tropical forms, now often segregated into several 
genera. Many arc insular and coastal forms, Schimper recording 4 
species in the Indomalayan strand flora and several species ranging 
eastward in the Pacific to the Fiji, Tonga, and Samoan Islands, and 
their fruits are recorded by both Gaudichaud and Guppy in the sea 
drift, although the oriental species are normally distributed by fruit 
])i.fireoiis (Mosley, Henisley. (iuppy). 

Do Candolle and Mi<|uel both considered the foliage, e.specially the 
^ enation, as offering the best criteria for diffeivntiation, but in the 
absence of comparative material and the incomplete character of the 
Panama fossil it is not possible to apply these criteria. The American 
Becent species number about 25, and these are mainly South Ameri- 
can in their distribution, although the sections or genera Virola 
Aublet and Compsoneura De Candolle both occur in Central America. 

The distribution of the Kecent species in tropical America, Asia, 
and Africa is conclusive evidence of a Tertiary history, although this 
evidence is practically unknown. Geyer ^ described two forms of leaf 
fragments from the Miocene of Labuan (Borneo) and Engelhardt* a 
third from the Tertiary of Ecuador and Chile. The most conclusive 
evidence of their Tertiary radiation is furnished by the characteristic 
fruits described recently by the writer* and preserved in the wind- 
blown sands of the uppermost Eocene of Texas. 

Occurrence. — Culebra formation (upper part). East wall of Gail- 
lard Cut just north of Canal Commission station 1760 (collected by 
M. I. Goldman). 

Order ROSALES. 

Superfamily LEGUMINOSAE, 

Genus TAENIOXYLON Felix. 

TAENIOXTLON MULTIRADIATUM Felix. 

Plntt?s 14 and 15. 
Tacnioxylon muUinidiatum Fklix, Die fossilen HOlzer We8tlndieii& 
Sanmil. pnlaoont. Al»li.. ser. 1, Hoft 1, p. 11, pi. 1, figs. 10, 11; pi. 2. 
flj?. 10, 1883. 

Transverse section. — ^In a radial distance of 5 cm. there are no 
defiJiite annual or seasonal rings. In certain zones the vessels are 

» Ccyl.r, II. T., Vcya Kxpedlllon, voL 4, p. 498, pi. 33, flgs. 3-6. 1887. 
< Engelhard t, U., Abb. Senck. Naturf. Gesellsch., toI. 16, p. 663, pi. 6, flsr. 9; pi. 7. 
flK. 12, 1891 ; vol. 19, p. 13, pi. 1, fig. 21, 1895. 

« BtM-ry. E. W., Amer. Journ. Scl., ser. 4, vol. 42, pp. 241-245, fijfs. 1-6, 1916. 



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GBOJuOGY AND PALEaNTOLOGY OF THE CANAL ZONE. 31 

larger, more generally compound, and closer together, and in other 
zones they are more distant, slightly smaller, and prevailingly single. 
No changes are observable in the other elements and there is no 
regular alteration of vessel rich and vessel poor areas nor any change 
from so-called summer to spring wood such as characterizes the trees 
of the temperate zone. 

Vessels single or two, three, or four together in radial rows (an 
anomalous group of five vessels in juxtaposition is shown in the de- 
tailed drawing). Outline of single vessels elliptical, those in groups 
flattened on one or both sides by mutual compression ; their tangen- 
tial diameter ranging from 0.10 mm. to 0.14 mm. ; their radial diam- 
eter ranging from 0.12 mm. to 0.16 mm., exceptionally large ones up 
to 0.22 mm. ; their walls thick, 0.0067 nmi. to 0.01 mm. in thickness, 
clearly sliowing the numerous small pits in section. Vessels fre- 
quently filled with gum. Vessels usually surrounded by one to three 
layers of rounded or more or less compressed thin-walled wood paren- 
chyma, somewhat variable in amount in different parts of the stem 
and tending to form tangential bands. Prosenchyma very abundant, 
the elements polygonal, small, somewhat smaller than those of the 
wood parenchyma, and thick walled. Bays very numerous, one or 
two cells wide as seen in transverse sections, flexuous in their courses 
since they are bowed out around the large vessels and approach more 
or less in the radial intervals between vessels; from 0.10 mm. to 
0.20 mm. apait, averaging nearer the former than the latter figure. 
The ray cells toward the ends of the rays which appear to be those 
usually seen in the several sections examined are not elongated radi- 
ally but are nearly isodiametric and about 0.02 mm. in diameter. 

Radial section. — ^The radial section shows the close set, fine, trans- 
versely elongated pits of the vessels which have simple perforations. 
The wood parenchyma is septate, the cells being about 3^ times as 
long as wide with large simple pits. The rays are of variable height, 
from 9 to 17 cells. They ai'e seen in radial view to consist of a central 
aeries of radially elongated cells with numerous fine simple pits, 
above and below which is a series of longitudinally elongated cells, 
beyond which are one or two rows of isodiametric cells which are 
regularly hexagonal in this view. 

Tangentud section. — The tangential section shows the uniform 
close set fine pitting on all the walls of the vessels, the relative short 
length and the large simple pits of the luijoining septate wood paren- 
chyma. The rays are seen to be very numerous, and separated by 
but few rows of flexuous prosenchyma; they are lenticular in outline 
and of variable height, one or two rays of terminal cells (those which 
are hexagonal in outline in the radial view) are single; then come 
one to three biseriate rows (those longitudinally elongated in the 



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32 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

radial view) ; toward the median region the rays are three or four 
cells broad (the radially elongated cells in the radial view). 

Felix states that in the Antigua material the rays were usuall}^ 
biseriate, while uniseriate and triseriate rays were rare. I do not 
know the extent of his material, but in the case of that from Panama 
I had but few radial sections cut. Ray cells frequently filled to a 
greater or less degree with gum. 

Remarks. — Fragments of the wood of this species are very com- 
mon in the collections from Panama, but a good deal was rather badly 
decayed before petrification. That which has formed the chief basis 
for the foregoing description and all of the photographs and draw- 
ings is beautifully preserved. The species is clearly identical with 
the type, as very insufficiently described and illustrated by Felix. 
One highly ferruginized and fairly well preserved quadrant of a 
trunk indicates a large tree, with a diameter of at least 25 cm. 

The genus Taenioxylon was established by Felix in 1882 with 
T. variams from Antigua as die type. He has since described 7 addi- 
tional species including 2 additional from Antigua, 1 from southern 
Brazil, 1 from East Indies, 1 from Philippines, 1 from Caucasus, 
and 1 from the Swabian Alps. All are of Tertiary age and show 
resemblances to various members of the 3 Leguminous families, 
Caesalpiniaceae, Miomosaceae, and Papilionaceae. Felix considers 
the present species to be a member of the Papilionaceae, and it agrees 
entirely with Solereders accoimt of the anat<Mny of this family. Tlie 
two kinds of ray cells described have, according to Saupe, been 
shown to occur in the following tribes in this family, namely the 
Podalyrieae, Genisteae, Galegeae, Hedysareae, and Sophoreae. With- 
out much recent comparative material, which is unavailable, it is 
impossible to allocate the present species more definitely within this 
extensive family. 

Occurrences. — Bohio formation, middle Bohio Ridge (poorly pre- 
served) quadrant of a large trunk indicating a tree with trunk at 
least 25 cm. in diameter. Cucuracha formation, upper part. Green 
clays of Gaillard Cut (locality 6845) Oligocene limestone. Orbi- 
toidal limestone, 2 miles north of David (locality 6523) (all above 
collected by D. F, MacDonald). Culebra formation, upper part. 
Near top of big slide, just north of Culebra. Collected by M. I. 
Goldman (figured material). 

GoUections. — ^U. S. National Museum, Johns Hopkins University. 

Family MIMOSACEAE. 
Genoa IN6A Willdenow. 

INGA OLIGOCAENICA, new species. 

Plate 16, fig. 2. 
Description. — ^Leaflets rather above medium size, elliptical-ovate 
and very inequilateral in general outline. Apex abruptly acute, not 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 33 

extended. Base very inequilateral, truncate or ascending on one 
side and wide and cordate on the other. Margins entire, full. Tex- 
ture subcoriaceous. Length about 8 cm. or 9 cm. Maximum width, 
at or slightly above the middle, about 4 cm. Petiolule curved, short 
and stout, about 3 mm. long. Midrib stout, greatly curved. Sec- 
ondaries thin, five or six ^irs, angles of divergence and courses 
various, all ultimately camptodrome; lower pair opposite, from the 
top of petiolule ; they diverge from the midrib at angles of about 45 
degrees, curving slightly outward and then ascending, parallel with 
the respective margins; the one in the narrow side of the lamina 
arches along the margin in a brochiodrome manner; the one in the 
wide side of the lamina sends oflf on the outside a series of regularly 
spaced camptodrome tertiaries. Tertiary venation for the most 
part obsolete. 

This characteristic species may be compared with Inga densifiora 
Bentham,* Inga edulis Martins,* Inga marginata Willdenow,* or Inga 
speciosa Spruce * and with various other of the larger-leafed species 
of Inga in the American Tropics to which region the 212 of its 
existing species of shrubs and trees are confined. It may also be 
compared with a number of tropical American species of Cassia^ as, 
for example, Cassia rusetfoUa Jacquin. 

About fiften fossil species have been referred to Inga. These 
include three from the Upper Cretaceous, two European, and one 
North American. There are also two or three species in the Oligo- 
cene of Europe, one in the Pliocene of Bolivia, two in the Tertiary 
of Ecuador, and one in the Tertiary of Colombia, four well-marked 
species in the Lower Eocene of the Mississippi embayment (Wilcox 
Group) and one in the middle Eocene of that region (Claiborne 
Group). The Panama species is not especially close to any of the 
foregoing. It is nearest, however, to Inga latifoUa^ described by 
Engelhardt ■ from the Tertiary of Ecuador, differing in its broader 
fonn and more inequilateral base. 

Pittier records 14 existing species of Inga^ from Panama.* Hems- 
ley lists 35 species in his flora of Central America, or which number 
18 are recorded from Panama. 

Occurrence. — Lower part of Culebra beds one-fourth mile south of 
Empire Bridge. (Collected by D. F. MacDonald.) U.S.G.S. 6887. 

Type.—C2X. No. 36311, U.S.N.M. 

^Bentham, Trans. Linn. Soc Lond., vol. 80, p. 617, 1875 (Peru). 

»Martlu«. Flora, toI. 20, Belbl., p. 113. 1837 (Braall). 

' WUldenow, 8p. PI.. toI. 4, p. 1015, 1806 (Venezuela). 

'Spruce, In Bentham, Trans. Linn. Soc. Lond.. vol. 30, p. 620 (Brazil). 

'Engelbardt, H., Abb. Senck. Natnrfor. Oesell., vol. 10, 1805. p. 20. pi. 2, flgs. 11, 12. 

•Pittler, H.. Cont. V. S. Natl. Herb., vol. 18. pt. R. pp. 218-223. 1016. 



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84 BULLBTIlfr 108, UNITED 8TATBS NATIONAL MUSEITM. 

Fanuly CAESALPINIACEAE. 
Cr^as CASSIA liimaaiis. 

CASSIA CULBBRBNSIS. imw spmIm. 

Plate 16, fig. 1^ 

Description, — Leaves obviously pinnately compound. Leaflets 
ovate, slightly inequilateral and falcate, with an obliquely aciuni- 
nate, practically equilateral tip, and an acuminate markedly inequi- 
lateral base. Length about 6.25 cm. Maximum width, about mid- 
way between the apex and the base, 2.75 cm. ; one side of the lamina 
15 mm. wide, the other 12.5 nmi. wide. Texture mediimily coriace- 
ous. Petiolule reduced to a thickened proximal part of the midrib 
extending but 1 mm. below the point of junction of one margin and 
about 2.5 mm. below the point of junction of the opposite margin. 
Margins entire, evenly rounded and full. Midrib relatively thin, not 
prominent, curved. Secondaries thin, numerous, about 10 suboppo- 
site to alternate pairs; they diverge from the midrib at wide angles, 
about 70° in the middle part of the leaflet, are nearly straight regu- 
larly spaced and subparallel in their outward course for two-thirds 
of the distance to the margin where the principal ones fork to join 
in rounded arches the similar branches of adjacent secondaries; the 
secondaries in the apical and basal portions of the leaflet are regu- 
larly camptodrome; those toward the tip of the leaflet more closely 
spaced. Marginal tertiaries camptodrome, internal tertiaries mostly 
obsolete. 

This type in its general form and the character of its base and 
petiolule indicates that it is a leaflet of a pinnate leguminous leaf. 
Its general appearance suggests comparisons with the genera 
Sweetiaj Myrocarptcs^ Toluifera^ Cassia^ and Sophora — the first three 
confined to tropical South America and the last two cosmopolitan in 
the existing flora. While the evidence is not conclusive, I prefer to 
consider it more closely allied to Cassia than to the other genera 
mentioned, particularly as the venation characters are such as I have 
considered referable to Cassia in my studies of the fossil floras of the 
southern United States. No species related to the Panama form is 
known from the Oligocene of the United States. 

The modern species of Cassia are very niunerous, upwards of 400 
having been described. They comprise herbs, shrubs, and trees of 
varied habitats in the warmer parts of both hemispheres, particularly 
tropical America. The fossil species are also numerous and the 
generic history goes back to near the base of the Upper Cretaceoua 
The genus has been continuously represented in the warmer parts of 



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OEOLOOY AND PALEONTOLOGY OF THE CANAL ZONE. 35 

America from the time of deposition of the Tuscaloosa sediments of 
Alabama to the present. 

Occurrence. — Culebra formation, lower part, one-fourth mile south 
of Empire Bridge (collected by D. F. MacDonald) U.S.G.S. 6837: 

Type.—CKt. No. 36312, U.S.N.M. 

Order GERANIALES. 

FamUy MALPlGHIACEAE. 
Genus HIRAEA JacquiiL 

mRAEA OUGOCABNICA, new ipMtos. 

Plate 17, fig. 1. 

Descriptuyn. — ^Leaves relatively large, ovate-lanceolate in outline, 
falcate, with an equally cuneatly pointed apex and base. Margins 
entire, evenly curved. Texture subcoriaceous. Length about 9.5 
cm. Maximum width, at or somewhat below the middle, about 3.5 
cm. Petiole short, stout, about 3 mm. in length. Midrib stout, flex- 
aous. Sec<mdaries thin, regularly spaced, about 9 pairs, prevailingly 
alternate; they diverge from the midrib at angles of about 45° and 
sweep upward in regular subparallel slight curves, and are campto- 
drome in the marginal region. Tertiaries obsolete. 

This genus, which has well characterized leaves, has seldom been 
recognized in the fossil state. One species ^ is not uncommon in the 
lower Eocene of the Mississippi embayment, and Ettingshausen ' 
has recorded, but not described, a second species from the Ypresian 
of Alum Bay, England. 

The existing species number between 25 and 30 and are exclusively 
American, ranging from Mexico and the Antilles throughout Central 
and northern South America to the Peruvian tropica 

The present fossil species is not unlike Hiraea wilcoxiana Berry * 
from the lower Eocene of Tennessee and is closely comparable with 
the existing Hiraea chrysophyUa Jussieu of the northern coastal re- 
gion of South America. 

Occuarence. — Caimito formation 7 miles northeast of Bejuca 
(U.S.G.S. station 6840). Collected by D. F. MacDonald. 

ryp^._Cat No. 85818, U.S.N.M. 

Genus BANISTERIA Linnaeus. 

BANISTEBIA PRABNUNTIA. new species. 

Plate 17. fig. 2. 
Description. — ^Leaves of medium size, broadly ovate in general out- 
line, with an abruptly acuminate tip and ajbroad rounded or cuneate 
hase. Length about 8 cm. Maximum width, at or slightly above the 

* Berry. E. W.. U. & Oeol. Surrey Prof. Pap«r 91. p. 257, pi. 57, fig. 8 ; pi. 109. fig. e. 

* Bmngshaiwen. C tod. Roy. Soc. London Proc.. Tol. 30. p. 285. 1880. 
*U. 8. Geol. Snrrey Prof. Paper 91. p. 257. 



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36 BULLETIN 103, UNITED STATES NATIONAL MUSBUM. 

middle, about 5 cm. Margins entire, full and rather evenly roimded. 
Petiolar character unknown. Midrib of medium size, uncharacteris- 
tic. Secondaries thin, seven or eight opposite to alternate pairs di- 
verge from the midrib at regular intervals at angles varying from 
45° in the upper part of the leaf to 56° in the basal part ; they ascend 
in slight but subparallel curves increasing in intensity as they pro- 
ceed toward the margins with which they become subparallel and 
eventually camptodrome. Tertiaries thin, mostly obsolete. Lieaf 
substance thin but apparently of a somewhat coriaceous texture. 

The present species receives its name from its supposed praenuntial 
relationship to the existing Banhteria mvemariensia De Candolle, a 
form ranging from the West Indies to Brazil and whose somewhat 
variable leaves may be exactly matched by the fossil. 

The genus contains upward of eighty existing species, mostly 
climbing shrubs, confined to the American tropics and largely de- 
veloped in northern South America. Its geological history goes back 
to the Lower Eocene, a species having been described by Watelet from 
the Ypresian of the Paris basin and four homotaxial species, one 
based on seeds, having been described by the writer from the Wilcox 
group of the Mississippi embayment in Western Tennessee and Ken- 
tucky. Several additional fossil species have been described from 
the European Tertiary, from all of which the Panama fossil is 
conspicuously different, its major differential character being its 
relatively short and broad outline. 

A species based upon fruits has been described by Engelhardt^ 
from the Tertiary of Ecuador. 

There are 5 species of Banisteria recorded by Hemsley from Cen- 
tral America, 3 of these in Panama, B. hiUhergiana Beurling on the 
seashore of the island of Manzaiiillo. Two additional Panama spe- 
cies of Banisteria are referred to the allied genus Heteropterya Kunth 
by Hemsley. 

Occurrence. — Culebra formation. West wall of Gaillard Cut be- 
low Miraflores locks (collected by M. I. Goldman). Culebra forma- 
tion (lower). West wall of Canal opposite Culebra Railroad station. 
(Collected by D. F. MacDonald). 

Family EUPHORBIACEAE. 
Genus HIERONYMIA AUem. 

HIERONTMIA LEHIIANNI Eng^lhardt (7). 

Plate 16, fig. 3. 
Hieronymia lehmanni Enghxhabdt, t)ber neue Tertiarpflauzen Sdd-Ameri- 
kas. Abh. Senck. Natuff. Gesell., vol. 19, p. 11, pi. 2, figs. 1, 2, 1895. 
Description, — ^Leaves broadly elliptical or somewhat deltoid and 
inequilateral in outline, with a shortly acuminate tip and broadly 

^ Eogelhardt, H.. (jber neue Tertiftrpflanxen Slld-Amerlkas, Abh. Senck. Natnrf. Ge- 
Pellsch., vol. 19, p. 14. pi. 2. figs. 18, 19, 1895. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 37 

rounded full lower lateral margins and a very wide, somewhat ob- 
liquely truncated base. Length about 12 cm. Maximum width, in the 
lower half of the leaf, about 10 cm. Margins entire, full, and rounded. 
Texture thin but coriaceous. Midrib stout, curved, prominent on the 
lower surface of the leaf. Secondaries stout, 10 or 11 irregularly 
spaced pairs, prominent on the lower surface of the leaf ; they diverge 
from the midrib at wide angles which become more acute in the apical 
part of the leaf, those on the narrower side are more ascending and 
somewhat straighter than those on the wide side, all are conspicu- 
ooflly camptodrome at some distance from the margin. Tertiaries 
thin, mostly percurrent. Areolation of small, isodiametric polygonal 
meshes, wdU marked on the under side of the leaf. 

This large leaf is unfortunately represented by fragmentary mate- 
rial from a single locality in the Caimito formation. In some 
respects its characters suggest a broad FicWj but it seems clearly 
identical with the species described by Engelhardt^ in 1895 from 
the Tertiary of Ecuador. I have, however, queried the determina- 
tion because of the broken character of the Panama material. In 
the illustration I have reconstructed a leaf from a combination of 
the Panama material with the more complete specimens figured 
by Engelhardt from Ecuador. The two largest fragments from 
Panama are indicated on the drawing by tinting. It is unfortunate 
for purposes of correlation that the present determination can not be 
•conclusive, although in view of other similarities shown between 
the Oligocene plants of Panama and those from the Tertiary of 
Ecuador, I am disposed to regard the present determination as 
fairly satisfactory. 

The genus Hieronymia comprises about a dozen existing species 
of shrubs and trees confined to tropical America and rather widely 
distributed from Mexico to Brazil as well as in the West Indies. 

Occurrence, — Caimito formation, 7 miles northeast of Bejuca 
(U.S.G.S. station No. 6840). (Collected by D. F. MacDonald.) 

Collection.— V. S. National Museum, Cat. No. 35814. 

Order SAPINDALES. 

Famfly SAPINDACEAE. 
Genus SCHMIDELIA Linnaeus. 

SCHBODELIA BEJUCBN8IS. new species. 

Plate 17, flg. 4. 
Description. — ^Leaf or leaflet elongate elliptic in outline, inequi- 
lateraL Apex and tip equally and bluntly pointed inequilateral. 
Margins entire. Texture coriaceous. Length about 11 cm. Maxi- 

1 Ober neue Terti&rpflanzen Sttd-Amerikas, toI. 19, p. 11, 1896. 



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38 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

mum width, midway between the apex and the base, about 4.5 cm. 
Width on one side of the midrib 21.5 mm., on opposite side 24 mm. 
Petiole missing. Midrib flexuous, stout, and prominent. Secondaries 
stout, regulariy spaced, mostly immersed, i^ut 7 alternate pairs 
diverge from the midrib at angles of about 60°, curving upward 
subparallel and camptodrome in the marginal region. Tertiaries 
mostly obsolete, a few percurrent ones seen. 

This large and striking leaf is referred to the sapindaceous genus 
Schmidelioj which comprises about 100 existing species of the equa- 
torial regions of both hemispheres with unifoliate or palmately com- 
pound leaves. About half of the species are American where thej 
are confined to the Antilles, Central, and tropical South America. 
They are sometimes referred to the genus AUophylvs Linnaeus (as 
by Badlkofer) and with the exception of this genus all of the mem- 
bers of the tribe Thouinieae are confined to America. Fossil repre- 
sentatives have been unknown except for the petrified wood from 
the Oligocene of the island of Antigua which Felix described as 
Schmideliopsis.* 

Occurrence. — Culebra formation. East wall of GaiUard Cut just 
north of station 1760 (collected by M. I. (Goldman). 

Caimito formation, 7 miles northeast of Bejuca (U.S.G.S. 6840). 
Collected by D. F. MacDonald.) 

Type.—C^it. No. 35315, U.S.N.M. 

Ordet THYMELEALES. 

Family LAURACEAE. 
Genim MESPILODAPHNE Neea. 

MBSPILODAPBNB CULEBRBNSia new bpmIm. 
Plate 17, fig. 8. 

DeBcriptian. — ^Leaves lanceolate-falcate in general outline, with 
acuminate apex and base. Margins entire. Texture subcoriaceous. 
Length about 10 cm. Maximum width, in the middle part of the 
leaf, about 2.5 cm. Petiole missing. Midrib stout, curved, prominent 
on the under surface of the leaf. Secondaries stout, remote, regu- 
larly spaced, nine or ten subopposite to alternate pairs, they diverge 
from the midrib at angles of about 65 degrees and are conspicuously 
camptodrome close to the margins. Tertiaries obscured by the poor 
preservation of the material. 

The present species resembles numerous existing and fossil species 
of Lauraceae, from all of which, however, it appears distinct. It is 
similar to MespUodapJvne colv/mhiana Berry of the Upper Claiborne 
of the Mississippi embayment, but is a stouter, more falcate, shorter, 
and less acuminate form. 

^ Felix, J.. Die fossile HSlser Westindlens. p. 16, pi. 2. figs. 6. 8. 1888. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 89 

The modem species of MespUodaphne are numerous, inhabiting 
Africa and tropical America, and are often united with Oreodaph/ne 
and Stn/chnodaphne to form the composite genus Ocotea of Aublet 
Their fossil history is almost entirely lost in the multitude of species 
that have been referred to the form genera Lauras and LaurophyJr 
him. MespUodaphne is abundant and varied throughout the Eocene 
and Oligocene of the Mississippi embayment area. 

Occurrence. — Culebra formation, upper part. East wall of the 
GaiUard Cut just north of Canal JZone station 1760. (Collected by 
M. I. Goldman.) 

Order MYRTALES. 

Family MYRTACEAE. 
Genoa CALYPTRANTHES Swartz. 

CALTPTRANTHBS GATUNENSIS, m 



Plate 18. fig. 1. 

Description. — Leaves broadly oblong-elliptic in general ojatline, 
widest in the middle and tapering equally in both directions to the 
abruptly acute apex and base. Margins entire. Texture subcoria- 
ceous. Length between 7 cm. and 8 cm. Maximum width between 
3.5 cm. and 4 cm. Petiole missing. Midrib stout, somewhat curved, 
prominent on the lower surface of the leaf. Secondaries thin, very 
numerous, and close set, often inosculating by forking; they diverge 
from the midrib at angles averaging about 70 degrees, at intervals 
of 1 mm. to 3 mm., pursue a but slightly curved outwardly ascending 
course and have their ends united by an aerodrome vein on each 
edge of the lamina parallel with and from 1 mm. to 2 mm. within 
the margin. Tertiaries forming open isodiametric polygonal meshes. 

The present well-marked species closely resembles the only other 
named fossil form Calyptranth^es eocenica Berry from the lower 
Eocene of the Mississippi embayment (Wilcox Group). It may also 
be compared with the slightly smaller Myrtus rectinervis described 
by Saporta^ from the Sannoisian of southeastern France. 

The genus CaLyptranthes^ which is exclusively American in the 
existing flora, has about seventy species ranging from Mexico and 
the West Indies to southern Brazil. There is a strong generic like- 
ness between the leaves of all of the species. Calyptranthes zyzygium 
De Candolle may be mentioned, among others, as a form with leaves 
almost exactly like the fossil. There is also a marked family resem- 
blance to some of the existing tropical American species of EugeniOy 
and more especially Myrcia^ Myrcia nmltiilora De Candolle from 
the Guianas being very similar to the present species. 

*8Aporta, Ifttodct, toI. 1, p. 251, pi. 11. flg. 5, 1868. 



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40 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Hemsley records 7 existing species of Calyptranthes from Central 
America, two of which occur in Panama. 

Occurrence. — Gatun formation, Gatmi Borrow Pits. (Collected 
by M. I. Goldman.) 

Family MELASTOMATACEAE. 
Genus MELASTOMITES Unger. 

MBLASTOIOTBS MICONIOmBS, new simcIm. 

Plate 18, fig. 2. 

Description, — ^Leaf oblong-elliptic in outline, of relatively small 
size, with an equally and bluntly pointed apex and base. Length 
about 6 cm. Maximum width, in the middle part, about 2.25 cm. 
Margins entire. Texture subcoriaceous. Petiole short and stont. 
Midrib stout and prominent. Lateral primaries stout, prominent, 
diverging from the midrib at an acute angle just above Uie base 
and aerodrome. From the disposition of the outwardly directed 
nervilles from the primaries it is probable that subordinate wsco- 
drome primaries constitute an infra marginal vein on each side, bat 
these can not be made out. Close-set subparallel nervilles run trans- 
versely between the midrib and the primaries. 

This species is represented by a small amount of fragmentary 
material, too poor to permit definite generic determination. It is, 
therefore, referred to thi form-genus McHastomites proposed by lin- 
ger for generically undeterminable leaves of the Melastomataceae. 
While the fossil somewhat suggests the leaves of various Lauraceous 
genera, such as Cinnamom/um^ Com/phororrvaea^ Goeppertia^ and 
Cryptocarya^ its characters are clearly those of the Melastomataceae. 
It particularly suggests the genus T^ouchina Aublet, which has up- 
ward of 200 species of shrubs and undershrubs in tropical America. 

The family Melastomataceae is a relatively large one, with about 
150 genera and over three thousand species. It is almost strictly 
tropical, although some members range southward to 40^ south lati- 
tude. This great family is typically American, seven of the fifteen 
tribes into which it is divided being confined to tropical America, 
and about 2,500 of the existing species being also endemic in this 
region. While the geologic history of this vast assemblage of forms 
is practically unknown, there is no evidence to disprove the theory 
that it, like the allied families Combretaceae and Myrtaceae, had its 
origin in that most prolific region — ^the American tropics. 

The few fossil forms that have been found, including leaves, flow- 
ers, and calices, have been referred to the form-genus Melastomites 
first proposed by Unger. A doubtfully determined species, which 
probably belongs to the Lauraceae, has bewi recorded from the Up- 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 41 

per Cretaceous of Westphalia. The only known Eocene species i& 
the well-marked form present in the lower Eocene of the Mississippi 
embayment region (Wilcox Group.) Four Oligocene species have 
been described from Bohemia, Styria, and Egypt; four Miocene 
species from Switzerland, Prussia, and Croatia; and a Pliocene 
species from Italy. 

Occurrence. — Culebra formation, upper part. East wall of Gail- 
lard Cut just north of Canal 2k>ne station 1760. (Collected by M. I. 
Goldman.) 

Order EBENALES. 

Family EBENACEAE. 
GeniiB DISOPYROS Unnaens. 

DI08PTB08 MAGDONALDI, new bimcIm. 
Plate 18, figs. 4-8. 

Description. — Globose berry-like fruits of small size and consider- 
able consistency, possibly preserved in an unripe state since the flesh 
is stringy and with a gi-eat many tannin cells. The great abundance 
of these fruits in the andesitic tuffs makes it seem more probable, 
however, that they are mature, particularly as some are greatly flat- 
t^ed. The numerous elongated pendulous seeds and the amount of 
vascular fibers in the flesh would tend to prevent much compression 
in a certain number of cases. Diameter 12 to 15 mm. Flesh hard, 
very tannif erous, and with numerous fibers. Seeds 8 to 10 in number, 
(Wong, elliptical, compressed, with a hard seed coat. The interior 
of the seeds is filled with amorphous silica and fails to show any 
structure. Seeds about 7.5 nmi. long, averaging 3 mm. high and 1 
mm. to 2 man. thick, very unequally developed, one to three usually 
more or less abortive. Peduncle not preserved, nor do any of the 
specimens show the calyx. 

These seeds are exc^ingly abundant and more or less perfectly 
silicified, the flesh being dark brown and the seeds white, making 
very striking objects. They are clearly referable to Diospyros and 
90 far as I know represent the only taiown petrified fruits of this 
genus, although the persistent calices are not uncommon as impres- 
sions from the Upper Cretaceous onward. The modem species have 
from 4 to 12 compressed seeds which tend to become less niunerous 
^th the increase in the fleshy part of the fruit, so that possibly these 
more consistent and prevailingly 10-seeded fossil fruits may represent 
an earlier stage in their evolution, although this seems doubtful 
since the calyx of a very large fruited form is known from the Upper 
Eocene of southwestern Texas. 



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42 BULLETIN 103y UNITED STATES NATIONAL MUSEUM. 

Diospyros is cosmopolitan in the existing flora with about 180 
species in the wanner regions of both hemispheres. Mostly Oriental* 
but not unconmion in the southern United States, Antilles, and from 
Mexico through tropical South America. Upward of 100 fossil spe- 
cies are known ranging in age from the Upper Cretaceous to the 
present. 

Occurrence. — Section near mouth of Tonosi River, in deposits of 
Eocene age (MacDonald). 

Type.—C2X. No. 86316, U.S.N.M. 

Order RUBIALES. 

Family RUBIACEAE. 
Genus RONDELETIA Plnmier. 

RONDELBTIA GOLDMANI. new spaciM. 

Plate 18. fig. 3. 

Description. — Leaves lanceolate in outline, somewhat falcate and 
inequilateral, with an equally acuminate apex and base. Length be- 
tween 12 cm. and 13 cm. Maximum width, midway between the apex 
and the base, about 3 cm., 13.5 mm. on the concave side and 15.5 mm. 
on the convex side. Margins entire. Texture coriaceous. Petiole 
short and stout, expanded proximad, about 5 mm. long. Midrib 
curved, stout, and prominent. Secondaries thin, numerous, suboppo- 
site to alternate, rather regularly spaced ; about 15 pairs diverge from 
the midrib at angles of about 45° and ascend in rather flat but regular 
and subparallel curves and are camptodrome in the marginal region. 
Tertiaries obsolete. 

This well-marked species is referred to the subfamily Cinchonoideaa 
and tribe Rondeletieae and seems to Indicate an Oligocene species of 
Rondeletia^ a genus of shrubs and trees confined to tropical America 
and not heretofore found fossil. Rondeletia has about 70 existing spe- 
cies, a few of which occur in northern South America, but the ma- 
jority are confined to the Antilles (45 species) and Central America 
(24 species).^ The present species may be compared with the exist- 
ing Rondeletia raoemosa Swartz of Jamaica, and with other Antillean 
and Central American forms. More remote comparisons may be 
made with certain species of Psychotria^ as, for example, Psychotria 
barbiflora De CandoUe of Brazil, and with the genus Tapiria Jus- 
sieu of the Anacardiaceae, a fossil species of which, Tapiria lanceo- 
latdj has been described by Engelhardt* from the Tertiary of Ecua- 

^ Brltton records 36 species from Cuba. Bull. Torrey Hot. Club, toI. 44, pp. 20-30, 1917. 
'Bngelhardt, H., tjber neue Tertittrpflanzeii Sdd-Amerlkas, Abh. Senck. Naturf. Qesell. 
▼ol. 19. p. 16, pi. 9, fig. 4. 1896. 



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OSOLOOY AND PALEONTOLOGY OF THE OAKAL ZONE. 43 

dor. Another fossil species somewhat resembling the Panama form 
is Oinch&nidium multinerve described by Ettingshausen ^ from the 
Tertiary of Priesen, Bohemia. 

Named in honor of Dr. Marcus L Goldman, who collected it while 
a Fellow at the Johns Hopkins University. 

Occurrence. — Gatmi formation, Gatun Borrow Pits. (Collected by 
IL L Goldman.) 

Geniui RUBL4CrrES Weber. 

BUBIACrrES IXORBOmSS, new specias. 
Plate 18, figs. 9-12. 

Description. — ^Fruit bilocular, indehiscent or tardily dehiscent, 
hgneous, capsular-like. Form a prolate spheroid 2.7 cm. long and 2 
em. in diameter. The surface roughened by small tuberculations and 
pits. Walls about 2 mm. thick. Median partition thin. Seeds one 
m each cell, suspended, elliptical in both transverse and longitudinal 
sections, compressed along the central partition. Surface striate. 
Endosperm not ruminating. One seed is more fully developed than 
the other. The larger is about 2 cm. long, 1.4 cm. wide and 9 mm. 
thick. 

This well marked form is unfortunately represented by but a single 
specimen which however shows most of the cavity occupied by the 
fruit, the two contained seeds partially petrified and the lignified wall 
and part of the partition. The accompanjring illustrations show the 
atemal appearance of the fruit (fig. 9) and a side view showing the 
relative development of the two seeds (fig. 10). Figure 12 shows a 
lignified end of the fruit with the median partition and figure 11 
19 a side view with the smaller seed in front and the larger forming 
the background. So far as I know nothing like it has previously 
been found fossil. 

There seems to be no question but that the present fruit represents 
some Oligocene species of Bubiaceae and it is consequently referred 
to the form-genus RubiaeiteB proposed by Weber, although probably 
not congeneric with the previously described fossil species of Bubia- 
cites. The fruits of this large family exhibit considerable variety 
being either capsular, achene-Iike or drupaceous. Without a much 
larger amoimt of recent comparative material than is available it is 
not possible to definitely fix the botanical relation of the present 
species which, however, appears to be referable to the tribe Ixoreae or 
the Psychotrieae. The specific name chosen suggests a resemblance 
to the fruits of Ixora Linnaeus, a genus with over 100 species of 

^ EttingBhaiwen, C tod. Die FowUe Flora des TertULr-Beckens yon BiUn, TheU 2, p. 208, 
Pl. M. llg. 5, ISOS. 

SSTO*— ISg— Bull. 108 i 



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44 BXJLLETIK 103^ XJKITED STATES NATIONAL MUSBUM. 

shrubs and small trees found in the tropics of both hemispheres but 
chiefly Asiatic 

Occurrence. — Oatun formation. Qatun Borrow Pits. (Collected 
by M. I. Goldman.) 

EXPLANATION OF PLATES. 

Plate 12. 

Palmowylon palmadtes (Sprengel) Stensel. Cncnracha formation. 
Fig. 1. Showing abundance of fibrovascnlar bundles and gum c^ls. X20. 

Plate 18. 

Fig. 1. Ficu8 culel)rensi8 Berry. Culebra formation, 

2. Chiatteria ctUebrensia Berry. Culebra formation. 

8. Myristioophyllum panamense Berry. Culebra formation. 

Plate 14. 

Taenioxylon mulHradiatum Felix. Culebra formation. 

Fig. 1. Transverse section. X25. 

2. Same. X200. 

Plate 15. 

Taenioxylon multiradiatuin Felix. Culebra formation. 
Fig. 1. Radial section. X200. 
2. Tangential section. X200. 

Plate 16. 

Fig. 1. OasMia culehreMi9 Berry. Culebra formation. 

2. Inga oligocaenioa Berry. Culebra formation. 

3. Hieronymia lehmanni Engelhardt (?). Caimlto formation. 

Plate 17. 

Fig. 1. Hiraea oligocaenica Berry. Caimlto formation. 

2. BanUteria praenutUia Berry. Culebra formation. 

3. MespUodaphne culebrenHa Berry. Culebra formation. 

4. BchmideUa hefucenHs Berry. Caimlto formation. 

Plate la 

. Fig. 1. Cdtyptrtrnthes gatunensis Berry. Qatun formation. 
2. MeUutomites miconioides Berry. Culebra formation. 

8. Rondeletia goldmani Berry. Qatun formation. 
4-8. Diosyproa macdonaldi Berry. Eocene (?). 

4. Showing abundance of fruits in tufte. 
5,7,8. Transverse median sections of fruits. 
6. Longitudinal median section of fruit 
9-12. Rubiacites iworeoidea Berry. Gatun formation. 

9. External appearance. 

10. Median longitudinal section showing imequally developed seeds. 

U. Side view of seeds. 

12. Lignified fragment showing end walls and partition. 



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BULLETIN 103 PL. 12 




PALMOXYLON PALMACITES (SPRENQEL) Stenzel. 
For explanation of plate see page 44. 



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U. 8. NATIONAL MUSEUM 



BULLETIN 103 PL. 13 






Fossil Dicotyledonous leaves. 

FOK EXPLANATION OP PLATS SBK PAOS 44. 



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U. S. NATIONAL MUSEUM 



BULLETIN 103 PL. 14 




Taenioxylon multiradiatum Felix. 

Fob explanation of platc see paoe 44. 



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U. 8. NATIONAL MUSEUM 










Taenioxylon multiradiatum Felix. 

For explanation of plate see paqe 44. 



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BULLETIN 103 PL. 




Fossil Dicotyledonous Leaves. 

For explanation of pi-ate see page 44. 



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BULLETIN 103 PL. 17 






Fossil Dicotyledonous Leaves. 

For explanat.on or plate see page 44. 



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U. 8. NATIONAL MUSEUM 



BULLETIN 103 PL. 18 




Fossil Leaves. Fruits, and Seeds. 

For explanation or plate see page 44. 



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INDEX. 



Page. 

Acrostichuni 10,18.22,18 

auream 23 

Allophylua ^— , 88 

ADona w 1», 28, 29 

S^abn * 28 

marcgraytl ^ 27 

reticulata 28 

Asimina 28, 29 

triloba 28. 20 

ATicennia 16, 22 

BactrU 16 

BaateterU 20, 8ft, 86 

UUberglana 86 

praenimtia 18,28,88,44 

itnemarlenala 86 

Calyptrantbea 21, 89, 40 

eocenica 89 

gatunensis 18, 28, 89, 44 

tjtjgixim 89 

Camphoromaea *. — 40 

Cassia 20, 84 

cnlebrensia 18. 22, 84, 44 

maeifolla 88 

Chamaedorea — . 16 

Clchonldinm mnltinerTe 48 

Clnnamomam 40 

CcMnpsoneora 80 

Conocarpua 16.22 

Creacentia 16 

Cryptocarya 40 

Dloapyros ....^ 21, 41 

macdonaldl 18, 28 41, 44 

Kndogenites palmadt<a-,..,«.,«,.^ 24 

Bnsenia ■„,-,„- 21 

Faacieolltes palmadtwi «,« 24 

Ftcoa 19, fit 

cnlebrensia 18, 22, 86, 44 

Jynx 27 

newtonensis .. 26 

Ooeppertia _,»,„„ — 40 

Onatteria 19, 20, 87, 29 

cnlebrensls 
dolicbo] 
grandlflom 
onregon 
Heteropterys - 
Hippomane — 

Hlraea 

chrj 

oligocaenlca 18, 28, 88, 44 

wilcoxiana 86 

HIeronymla 17, 20, 86, 87 

lehmannl 18, 28. 86, 44 

Inga 20, 88,88 

densillora 88 

ednlli 88 




Paga. 
Inga latifolla 38 

oligocaenlca 18, 22, 82, 44 

Ixora 43 

Lagnncnlaria 22 

Laams 39 

Laurophyllnm :iu 

Melastomltes 40 

mlconioides 18, 28, 40, 44 

Melodomm 28 

Mespilodaphne 88, 30 

colnmbiana 88 

cnlebrensls 18,23.88,41 

Hltrephora 28 

Myrcia 21, 39 

mnltlflora 89 

Myrlstlca 20. 30 

Myrlstlcophylluni 20,29 

panamense-. 18, 22, 89, 44 

Myrocarpns 84 

Myrtns rectlnerrls 89 

Ocotea 89 

Oreodaphne 39 

Palmadtes dublni 24 

tenemm 24 

Palmozylon 17, «4, 25 

antlguense 25 

aschersoni 25 

ceylanlcnm 25 

Integmm 25 

mlBsisslppiensls 25 

palmadtes 18, 22, 84, 44 

•tellatnm 25 

tenemm 25 

▼arlabile 25 

Palms 16, 18, 23. 84 

Paritium 16 

Psychotrla barblflora.^ 42 

Rhisophora 16, 22 

BondeleUa 21, 48 

goldmanni 18, 23, 48, 44 

racemosa 42 

Rubiadtes 21, 48 

Ixoreoides 18, 23, 48, 44 

Sdimidelia 20, 87, 38 

bejncensis 18, 23, 87, 44 

Sophora 34 

SweetU 34 

Taenlozylon 17, 20, 80, 82 

mnltiradUtnm — 18, 22, 80, 44 

▼ariani 81 

Tapirla lanceolata 42 

Tlbonchlna 40 

Toluifera 34 

Trithrinax 16 

Venericardla planicosta 17 

Virola 80 

I 



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SMTTHSONIAN mSTITirnON 

UNITED STATES NATIONAL MUSEUM 

Bulletin 103 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



THE SMALLER FOSSIL FORAMINIFERA 
OF THE PANAMA CANAL ZONE 



By JOSEPH AUGUSTINE CUSHMAN 
Of the United States Geological Survey 



Extract from Bulletio 103, pages 45-87, with Plates 19-33 




WASHINOTON 

GOVERNMENT PRINTING OFHCB 

1918 



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THE SMALLER FOSSIL FORAMINIFERA OF THE 
PANAMA CANAL ZONE. 



By Joseph Augustine Cushman, 
Of the United States Geological Survey. 



INTRODUCTION. 

The collection of fossil foraminifera included in this report were 
sent to the writer by the United States Geological Survey. It con- 
sists almost entirely of material collected by Messrs. D. F. Mac- 
Donald and T. Wayland Vaughan in 1911, to whom I am indebted 
for data as to the geological correlation. The names applied to the 
geologic formations are those used in MacDonald's "Sedimentary 
formations of the Panama Canal Zone, with special reference to the 
stratigraphic relations of the fossiliferous beds," which appears in 
the latter part of this volume. Where former correlation has seemed 
not to apply to the foraminifera, especially those of three stations, 
6033(?, 6035, and 6036a, discussion of the data obtained from the 
foraminifera is given in detail later. 

The orbitoids and nimmiulites are both well represented in the 
collection, but as these require special study in connection with those 
of the Cioastal Plain and of the West Indian region it seems ad- 
visable to treat them in a separate paper which immediately follows 
the present one. 

The following data are given for only the stations from which 
foraminifera were obtained and which are recorded in this paper. 

LIST OF MATERIAL. 

OjS.OjS. BtaUon 6009. — OUgocene — Culehra formation (upper part) . 
From section in Canal cut 600 feet south of Miraflores Locks. 
Dark, soft, fairly well laminated clay rock. 
Few foraminifera and rather poorly preserved. 
6010. — OUgocene — CvZehra formation (lower part). 

From section — ^Pedro Miguel Locks to Paraiso Bridge. 
Dark, well laminated, very soft, carbonaceous clay rocks. 
Foraminifera in fairly good numbers and a rather varied assort- 
ment; mostly stained black, except certain of the Miliolidae, 
which still keep their calcareous tests moi-e or less in their 
original condition. 

46 



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46 BULLETIN 103, UNEDED STATES KATIOKAL MX7SEUM. 

6012. — OUffocene — CtUebra formation. 

From section — ^west side of Gaillard Cut 

a. Dark, well laminated soft and very friable carbonaceous shale. 

Few foraminifera — some glauconitic, others well preserved. 

c. From a lens of sandy limestone 5 feet thick. 

Few foraminifera — some stained, some glauconitic, rather poorly 
preserved as to details. 

d. From lenses of limy sandstone at base of gravel, 3 feet thick. 
Few foraminifera and these poorly preserved. 

6016. — OUffocene — Emperador Umeatone. 

From old quarry, one-fourth mile north of west from Empire. 

Cream-colored, coral limestone. 

Few foraminifera. 
6016. — OUffocene — Emperador Umestone. 

From old quarry, one-third mile north of west of Empire. 

Few poorly preserved foraminifera. 
6019. — Section on west side of Gaillard Cut near Las Cascadas. 

Or-f. OUffocene — Cvlebra fomuxtion. 

a. Grayish, rather nodular, impure limestone. 

Foraminifera few and poor. 

&. Dark, well stratified, very friable, tufaceous material. 

Foraminifera few and poor except Orhitolites^ which are large 
and fine. 

c. Grayish, well stratified, very friable, tufaceous sandstone. 
Few casts of foraminifera and central portions of orbitoids. 

d. Grayish-green, limy, tufaceous sandstone. 
Very few foraminifera, poor specimens. 

e. Thin-bedded, light gray to cream-colored, limy sandstone with 
some partings of light-colored clay. 

orbitoids and Orbitolitesf only. 

/. Dark, very friable shales and tuffs. 

Foraminifera fairly common, some well preserved, others glau- 
conitic. 

ff. OUffocene — Emperador Um^tone, 

Light gray to yellowish gray, somewhat sandy limestone. 

Some orbitoids and Orbitolitesf but little else in the way of 
foraminifera. 
60i0. — OUffocene — Cvlebra formation^ 

Same locality as 6019. 

Or^. Dark-gray carbonaceous clays, friable shales and tuffs. 

a. Foraminifera numerous but of few species, mostly glauconitic, 
at least in part. 

c. A few OrbitoUtes in the coralliferous layer. 



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QEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 47 

60JH. — Section in raikoay cuts near New Frijolea. 
a. Oligocene — Culehra formation. 
Dark, basic, orbitoidal, tufaceous material. 
Many worn central portions of Orbitoids and a very few other 
foraminifera poorly preserved. 
60£S. — Oligocene — Cvlehra formation (upper part). 

About 200 yards south of southern end of switch at Bohio 

Kidge station relocated line Panama Kailroad. 
Contains a number of species of foraminifera but for the most 
part broken or poorly preserved. 
6026. — Two miles south of Monte Lirio. 
Somewhat coarse-grained sandstone. 
Few poor specimens of foraminifera. 
60S9. — Section one-half mile from Camp Cotton^ toward Monte Lirio^ 
at big curve on railroad. Miocene — Gatun formation. 
a. Bluish, fossiliferous argillite. 
Very few foraminifera. 
h. Bluish argillite. 

Few foraminifera, but considerably more than in a. 
c. Bluish, fossiliferous argillite. 
Very few poor specimens of Amphistegina. 
6030' — Railroad cut north side of Big Swamp j one and one-half miles 
north of Monte Lirio. Miocene — Gatun formalion. 
Bluish gray, argillaceous beds. 

The only foraminifera consisted of a single specimen of TrUo^ 
ctUina. 
6031. — Section in cut one-half mile west of Camp Cotton toward 
Galun. Miocene — Gatun forTnaUon. 

Conglomerate bed and sandy marl 1 foot above. 
A few poorly preserved specimens of Quinqueloculina were the 
only foraminifera. 
6033. — Generalized section of the hVaffs exposed along the Pana/ma 
Railroad^ relocated line, about 3^500 feet south of Gatun Railroad 
Station. Miocene Gatun— rformation. 

c. Dark-colored, marly, fossiliferous clay. 
Rich in foraminifera, especially in specimens. A fair number 
of species, well preserved. 
6036. — Vicinity of Mindi Hill. Miocene — Gatun formation. 
Gray-green, fine grained sandy shell marl. 
Very fine-grained material, but with numerous species and speci- 
mens of foraminifera representing an off-shore assemblage. 
6036. — Monkey Hill, Mount Hope Station. Miocene — Gatun for- 
mation. 

Dark-colored, fine grained, sandy clay marl. 



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48 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Specimens of foraminifera numerous and well preserved, rep- 
resenting an off-shore assemblage comparable to 6085. 
6860. — Near Movm,t Hope — Pleistocene. 

Loose shells and marl obtained from ditch through swampy 
groimd about one-fourth mile from present sea beach and 
about 6 to 8 feet above high tide. 
Contains a few foraminifera of common shallow water, tropical 
species. 
. The geological position of certain material from near the Atlantic 
end of the canal seems from the evidence of the contained f oramin- 
fera to be younger than the position previously assigned to it — ^the 
upper Oligocene. By a reference to the table of distribution it will 
be noted that the great majority of the species occurring at the sta- 
tions in question; 6533o, 6035, and 6036, do not occur in the ma- 
terial of definitely Oligocene age. In such cases as that of Oristel- 
Icpria rotulata there is a slight difference in the specimens from these 
stations and those from the Pacific side, 6010, 6012a, 6012t?, but the 
specimens at the latter stations were in small quantity, and the dif- 
ferences could not be made use of, mainly from lack of a sufficient 
number of specimens. In the case of CristeUaria vaugKam this 
seems to be a well-characterized species occurring at several stations, 
but even in it there are very minor differences. Among the species 
of Glohigerina^ the more generalized species such as G. bulloideSj 
which has a very wide geological range, occur more or less constantly 
throughout the collections, but the strongest evidence comes from 
the last three species and Orbulina^ which are very rarely found 
fossil, and then only in the very latest tertiary. These were well 
characterized species, the specimens are very clean and complete, and 
resemble a modem Globigerina ooze of considerable depth. The 
three species of Pul/vinulina also occur nowhere but at these stations. 
PulvinuUna concentrica is essentially a recent species and P. me- 
nardii is characteristic of modem Globigerina ooze. Sigmoilina 
tenus and S. aspenda are also speces of recent Globigerina ooze of 
moderate depths. On the other hand, the lack of certain things is 
also significant. Amphistegina^ which occurs more or less regularly 
in the other portion of the material, is entirely wanting in the three 
Pacific stations, 6033(?, 6035, and 6036. Polystomella also does not 
occur. Both the last two genera are very characteristic of the 
coastal plain Oligocene of the United States. It may be argued in 
this case, however, that the stations were originally too far from 
shore to have these genera which are more characteristic of shallow 
littoral conditions. 

On the whole, the foraminifera b^r out the geological determina- 
tions based upon the other groups of organisms. 



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QEOUMIT AND PAI^OKIOIXMIT OF THE OAKAL ZONE. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONB. 51 

DESCRIPTIONS OF SPECIES. 

Family TEXTULARIIDAE. 
Genus TEXTULARIA Defrance, 1824. 

TBXTULARIA ABBRBVIATA iTOKWKny. 

Plate 19, fig. 1. 

TextuJaria abbreviata d'Orbigny, Foram. Foss. Bass. Tert. Vienne, 1846, 
p. 249, pL 15, figs. 9-12 (7-12). 

Description, — ^Test broad and short, somewhat compressed, cham- 
bers comparatively few in number, broad near the center and taper- 
ing to the periphery, sutures in these specimens indistinct, aperture 
an arched slit extending nearly across the test, wall comparatively 
smooth. 

Length 0.65 mm., breadth about 1 mm. Cat. No. 324608, U.S.N.M. 

Specimens from U.S.G.S. No. 6010, from the Culebra formation, 
dark clay north of Pedro Miguel Locks. Apparently the material 
is rather metamorphosed and more or less glauconitic so that little 
of the original test is preserved. This is a rather common Tertiary 
species. 

TEXTULARIA SAGITTULA Defrance. 

Plate 19, fig. 2. 

Temtvlaria sagUtula Defbance, Diet. Sc4. Nat., vol. 32, 1824, p. 177 ; vol. 53, 
1828, p. 344 ; Atlas, Conch., pi. 13, fig. 5. 

Description. — ^Test elongate, tapering, much compressed especially 
it sides, chambers numerous, sutures indistinct, aperture a curved 
dit occupying about one-half the width of the base of the chamber. 

Length about 1.5 mm., breadth 1 mm. Cat. No. 324609, U.S.N.M. 

A few poorly preserved specimens from U.S.G.S. No. 6025, from 
the Culebra formation, foraminiferal marl and coarse sandstone 
about 200 yards south of southern end of switch at Bohio Ridge 
station, relocated line, Panama Railroad. 

Altiiough this material is more or less glauconitic and poorly pre- 
served the three specimens, one of which is here figured, are referred 
wiUi a reasonable degree of certainty to this species. 

A single fragmentary specimen from U.S.G.S. No. 6026, from the 
Culebra formation, coarse, sandy foraminiferal marl about half way 
between Monte Lirio and Bohio Ridge, relocated line, Panama Rail- 
road, seems also to be this species. 



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52 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

TEXTULABIA AGGLUTINAN8 rOrUcnj. 

Plate 19, fig. 3. 

Textularia aggluUnans d*Okbiony, in De la Sagra, Hist Fis. Pol. Nat. 
Cuba, 1839, " Foramlnifdres," p. 136, pi. 1, figs. 17, 18, 82-34. 

Description. — Test elongate, tapering, but slightly compressed lat- 
erally, chambers high, sutures deep, outline sinuous, end view broadly 
elliptical, wall composed of rather coarse agglutinated material, 
aperture a narrow slit a little more than half the width of the base 
of the chamber. 

Length 1.28 mm., breadth 0.65 mm. Cat. No. 824610, U.S.N.M. 

A single specimen here figured seems referable to this species. It 
is from U.S.G.S. No. 6019-/, from the uppermost bed of the Culebra 
formation, the lower limestone of the Las Cascadas section, opposite 
Las Cascadas, Ghtillard Cut. Although not so rounded in end view 
as this species usually is in recent specimens, the general characters, 
wall structure, high rotimd chambers and lobulated outline seem to 
place it here. 

TEXTULARIA LAMINATA, new fpedM. 

Plate 19, fig. 4. 

Description. — ^Test elongate, cuneate, tapering from the widest 
part near the apertural end, gradually and evenly to the initial end 
which is subacute, median line raised thence tapering rapidly toward 
the periphery which is thin and extends out into a lamella-like 
border, chambers numerous, wide and low, sutural lines raised, some- 
what curved backward; border irregular, wall finely arenaceous; 
aperture indistinct. 

Length 2 mm., breadth 1.2 mm. 

Specimen figured from U.S.G.S. No. 6010, from lower part of the 
Culebra formation, dark clay north of Pedro Miguel Locks. Speci- 
men rather better preserved than most from this station. The end 
view of this specimen is mainly rhomboidal with the bordering 
carina extending outward in a thin carina. It is in some ways sug- 
gestive of Textularia carinata but differs in many respects from 
that species which is also figured on plate 19, fig. 6. 

Type'Specimen.—C9it. No. 324611, U.S.NJML 

TEXTULARIA SUBAGGLUTINANS, newspMlM. 
Plate 19, fig. 5. 

Description. — ^Test subrhomboidal in front view tapering from the 
middle toward either end, in end view oblong, sides tnmcated ; cham- 
bers comparatively few, somewhat inflated, sutures conspicuously de- 



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QSOLOOY AND PALEONTOLOaY 07 THE CANAL ZONE. 53 

pressed, wall composed of rather coarse arenaceous material, aper- 
ture extending into the base of the chamber in a narrow rounded 
opening deeper than wide. 

Length 1.3 mm., breadth 0.85 mm. 

This species was fairly common from U.S.G.S. No. 6033c, the 
Gatun formation, in marl from second bed from bottom, just below 
lower clay, Gatun section, relocated line Panama Railroad. 

This species may be distinguished from Textularia aggtlutirums by 
the truncated sides, the oblong end view and especially by the deep, 
narrow aperture. 

Type specimen.— Cnt. No. 324612, U.S.N.M. 

TEXTULARIA CARINATA d'Orbigny. 

Plate 19, fig. 6. 

Textularia carinata D'ORniGNV, Ann. Sci. Nat., vol. 7, 1826. p. 263, No. 23; 
Foram Foss. Bass. Tert. Vienne, 1846, p. 247, pi. 14. figs. 32-34. 

Description. — ^Test much compressed, rather abruptly tapering 
toward the initial end, sutures strongly limbate, in well-preserved 
specimens extending out from the periphery in angular spine-like 
projections, aperture narrow, elongate. 

Length 1 mm., breadth 0.65 mm. Cat No. 324613, U.S.N.M. 

The only material of this species is from U.S.G.S. No. 6036, from 
the Gratun formation, a dark-colored, fine-grained, sandy clay marl 
from Monkey Hill, Moimt Hope Station. It is very evidently this 
species and is well preserved. 

TEXTULARIA PANAMBN8I8, new tpeciea. 

Plate 20, fig. 1. 

Description. — ^Test rhomboid in front view, very much compressed, 
in end view long and narrow, the faces nearly parallel, sides rounded ; 
composed of comparatively few chambers but variable; long and 
low, sutures somewhat depressed, wall rather coarsely arenaceous; 
aperture indistinct. 

Length 0.85 mm., breadth 0.65 nmi. 

The figured specimen is from U.S.G.S. No. 6036, from the Gatun 
formation, a dark-colored, fine-grained sandy clay marl from Monkey 
Hill, Mount Hope Station. Specimens were common from U.S.G.S. 
No. 6033(7, in marl from second bed from bottom, just below lower 
day, Gatun section, relocated Panama Railroad. 

This is a rather striking species, with its very flat, broad front 
view and very compressed character of the test. 

Type-specimen.— Cst, No. 324614, U.S.N.M. 



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54 BULLETIN 103, UNITED STATES NATIONAL MUSBTTM. 

GenuB CHRYSAUDINA d'OtUgny, 1846. 

CHRYSALIDINA PULCHELLA, new fp«ciet. 

Plate 20, fig. 2. 

Description. — Test elongate, gently tapering, broadest at the apical 
end; in end view triangular; early chambers triserial, later ones uni- 
serial ; chambers in uniserial portion triangular, the sutures distinct, 
gently curved backward at the angles, outline more or less irregular, 
apertural face gently convex, with indications of numerous circular 
apertural openings, wall smooth. 

Length 0.5 mm., breadth 0.2 mm. 

This species occurred at U.S.G.S. No. 6036, the Gatun formation, 
in dark-colored, fine-grained, sandy clay marl, from Monkey Hill, 
Mount Hope Station. 

The species differs from the only known recent species, Chrysa- 
lidina dimorpha^ in the more tapering and elongate test, the greater 
irregularity of the contour and test in general and its generally less 
trim and neat appearance. The specimen figured is well preserved 
in its general characters, except those of the apertural face,- which 
are somewhat obscured. 

Type-8pecivi€n.—Cs^t. No. 324615, U.S.N.M. 

Genus BOLIVINA d'Orbigny, 1826. 

BOLTVINA cf. B. PUNCTATA d'Orbiffny. 

Plate 21, fig. 3. 
Bolivina punctata d'Orbigny, Voyage Am6r. M^rld., vol. 5, pt. 5, " Foraml- 
nif^res," 1839, p. 63, pi. 8, figs. 10-12.— H. B. Brady. Rep. Voy. Chal- 
lenger, Zoology, vol. 9, 1884, p. 417, pi. 52, figs. 18, 19.— Flint, Ann. 
Rep. U. S. Nat. Mtis., 1897 (1899), p. 292, pi. 38, flg. 1. 

Description. — Test much elongate, sides nearly parallel, abruptly 
tapering at the initial end, chambers numerous, usually higher than 
broad, inflated, sutures distinct but slightly depressed; wall finely 
punctate, occasionally becoming slightly striate. 

Length 0.60 mm., breadth 0.15 mm. Cat. No. 324616a, 6, U.S.N.M. 

Specimens which seem referable to this species were obtained at 
U.S.G.S. No. 6033c, Gatun formation, marl from second bed from 
bottom, just below lower clay, Gatun section, relocated line Panama 
Railroad and 6035, Gatun formation, from gray green, fine grained, 
sandy shell marl, vicinity of Mindi Hill. There is a tendency for 
the specimens to take on a semi-striate appearance, an extreme form 
both in shape and striation shown in plate 21, figure 3. 

BOLFVINA AENABIENSIS (Gotta). 
Plate 21, fig. 2. 
Brizalina aenarien^is Costa, Atti Acad. Pontanlana, vol. 7, 1856, p. 297, pi. 

15, flg. 1, A. B. 
Bolivina aenariensis H. B. Brady, Free Roy. Soc. Edinburgh, vol. 11, 1882, 
p. 711; Rep. Voy. Challenger, Zoology, vol. 9, 1884, p. 423, pi. 58, figs. 
10, 11. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 55 

Description. — Test much compressed, composed of numerous cham- 
bers about twice as broad as high, sutures distinct, slightly curved 
backward, chambers slightly inflated, especially in the center, test 
bordered by a narrow but distinct carina ; surface smooth except for 
several longitudinal raised costae radiating from the initial end which 
carries also a short spine. 

Length 0.65 mm., breadth 0.35 mm. Cat. No. 324617a, &, U.S.N.M. 

A few specimens were obtained from U.S.G.S. No. 6033c?, Gatun 
formatito, in marl from second bed from bottom, just below lower 
clay, Gratun section, relocated line, Panama Railroad. 

While these specimens are not absolutely typical they undoubtedly 
belong to this species. 

Very typical specimens occur at U.S.G.S. No. 6086, Gatun forma- 
tion, in dark colored, fine grained, sandy clay marl, from Monkey 
Hill, Mount Hope Station. 

BOUVINA ROBUSTA H. B. Bradr. 

Plate 21, fig. 4. 

Bolivina rohusta H. B. Brady, Quart. Joum. Micr. Sci., vol. 21, 1881, p. 
67; Rep. Voy. Challenger, Zoology, vol. 9, 1884, p. 421, pi. 53, figs. 7-9. 

Description. — ^Test compressed, gradually tapering toward the 
apical end; chambers comparatively few; about twice as broad as 
high; sutures limbate, gently curved backward, often slightly lobu- 
iated or occasionally showing traces of reticulation on the surface, 
wall otherwise smooth but punctate, not spinose at the apical end. 

Length 0.45 mm., breadth 0.25 mm. Cat. No. 324618, U.S.N.M. 

These specimens, an extreme form of which is figured, are many of 
them very close to typical B. robusta which is at best either a variable 
species or one including more than one form. The sutures are 
usually limbate, as shown in some of Brady's figures, but no apical 
spine is apparently in any of the specimens in this material. They 
were from U.S.G.S. No. 6035, Gatun formation, from gray green, 
fine grained, sandy shell marl, vicinity of Mindi Hill. 

BOLIVINA* ipMlMr 

Plate 21, fig. 1. 

This specimen is rather ill-defined and cannot be definitely deter- 
mined from the single example, the sutures are limbate as in Bo- 
Uvina rohusta Brady, but have apparently no secondary extensions 
as in that species. The whole specimen seems to be replaced. The 
specimen is from U.S.G.S. 6010, lower part of the Culebra formation, 
frwn dark clay north of Pedro Miguel Locks. Cat. No. 824619, 
U.&N.M. 



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56 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

GeniiB BI6ENERINA d'OrUsny, 1826. 

BIGBNBRINA NODOSABIA d'OrMgny. 

PlaU 21. fig. 5. 
Bigenerina nodosaria d'Obbionv, Ann. Sci. Nat, vol. 7, 1826, p. 261, pi. 
11, figs. 9-11.-— H. B. Bbady, Rep. Voy. Challenger, Zoology, vol. 9, 
1884. p. 869, pi. 44. figs. 14-18. 

Description, — ^Test elongate, subcylindrical, early portion consist- 
ing of a few chambers arranged as in TexttdariOy later ones uniserial, 
early portion tapering abruptly toward the apical end, wall coarsely 
arenaceous, sutures rather indistinct, aperture circular and centraL 

Length 2 mm., breadth 0.8 mm. Cat. No. 324620, U.S.N.M. 

Several specimens in excellent condition were obtained from 
U.S.G.S. No. 6036, Gatun formation, in dark-colored, fine-grained, 
sandy clay marl from Monkey Hill, Mount Hope Station. 

These specimens, as in the one figured, have but a slight indication 
of the biserial chambers from the exterior, but otherwise seem to be 
typical. At first glance they might be taken for a species of 
Clavulina. 

• Genus GAUDRYINA d'OrWgny, 1839. 

GAUDRTINA FUNTII CoBhmmn. 

Plate 20. fig. 4. 
Oaudryina suhrotundata Flint (not G. subrotundata Schwager, 1806), 

Ann. Rep. U. S. Nat Mus., 1897 (1899). p. 287. pi. 33. fig. 1. 
Oaudryina flintii Cushman, Bull. 71. U. S. Nat Mus., pt 2. 1911, p. 63, 

fig. 102(j-c. 

Description. — Test subcorneal, early portion rounded conical, trise- 
rial, later portion subcylindrical, biserial chambers of later portion 
nearly semicircular in transverse section, sutures distinct; wall are- 
naceous; aperture subcircular, at the base of the inner margin of the 
chamber. 

Length 1.20 mm., breadth 0.72 mm. Cat. No. 824621. 

A single specimen which seems to be close to recent specimens of 
this species was obtained from U.S.G.S. No. 6010, lower part of the 
Culebra formation, in dark clay, north of Pedro Miguel Locks. The 
specimen is somewhat glauconitic and certain of the details are more 
or less obscured. 

GAUDRYINA TRIANGULARIS Cluhmmn. 

Plate 20, fig. 3. 

Oaudryina triangularis Oushman, Bull. 71, U. S. Nat Mus.. pt. 2, 1911, 
p. 65, figs. 104a-<?. 

Description. — ^Test somewhat longer than broad, early portion tri- 
angular, the faces somewhat concave, triserial; later portion biserial, 



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QEOLOOY AND PALEONTOLOGY OP THE OANAL ZONE. 67 

rounded in transverse section; wall coarsely arenaceous, chambers 
comparatively few, sutures indistinct, aperture a narrow slit at the 
base of the inner margin of the last formed chamber. 

Length 1.7 mm., breadth 1.0 mm. Cat No. 324622, U.S.N.M. 

A single specimen which seems to belong to this species was found 
in material from U.S.G.S. No. 6010, lower part of the Culebra forma- 
tion, in dark clay, north of Pedro Miguel Locks. The specimen, like 
many others from this station, is glauconitic and not well preserved 
in all its details. 

Genus CLAVUUNA d'Orbigny, 1826. 

CLAVULINA PABISIENSIS d'Orbigny. 

Plate 20, fig. 5. 

OlawUna pariaiensia d'Orbigny, Ann. Sci. Nat., vol. 7, 1826. p. 268. — H. B. 
Bbadt, Rep. Voy. Challenger^ Zoology, vol. 9, 1884, p. 395, pi. 48, figs. 
14-18. 

Description. — ^Test elongate, subcylindrical, early portion conical, 
later portion gradually increasing in diameter toward the apertural 
Old, chambers comparatively few, those of the uniserial portion cir- 
cular in cross section, waU coarsely arenaceous, somewhat rough; 
aperture circular, terminal. 

Length nearly 2 mm., diameter 0.7 mm. Cat. No. 324623, U.S.N.M. 

A single specimen representing this species was obtained in mate- 
rial from U.S.G.S. No. 6010, lower part of the Culebra formation, 
in dark clay north of Pedro Miguel Locks. Both this and the fol- 
lowing are conmion Tertiary species. 

CLAVULINA COMinTNIS d'Orbigny. 

Plate 20, fig. 6. 

ClavuUna communis d'Orbigny, Ann. Scl. Nat., vol. 7, 1826, p. 268 ; Foram. 
Fobs. Bass. Tert. Vlenne, 1846. p. 196, pi. 12, figs. 1, 2. 

Description. — ^Test very elongate, subcylindrical, circular in trans- 
verse section, early portion triserial, later portion uniserial, of rather 
uniform diameter, sutures more or less indistinct, wall smooth ; aper- 
ture terminal. 

Length 2 mm., breadth 0.45 mm. Cat. No. 324624, U.S.N.M. 

A single specimen of this species occurred with the preceding, 
U.S.G.S. No. 6010, in the lower part of the Culebra formation. It 
is fragmentary but probably represents this species. 



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58 BULLETIN 103^ UNITED STATES NATIONAL MUSEUM. 

Genus YIRGULINA d'Orbigny, 1826. 

VIRGULINA SQUAMOSA d*0rblfn7. 

Plate 21, fig. 6. 

VirgiUma squamosa d'Obbiony, Ann. Sci. Nat., vol. 7, 1826, p. 267. — Gush- 
man, BuU. n, U. S. Nat Mus., pt 2, 1911, p. 91, fig. 145a, h. 

Description. — ^Test elongate, tapering gradually to the apical end 
and again toward the apertural end, chambers comparatively few, 
inflated, sutures distinct, wall smooth, aperture a comma-like slit at 
the base of the last formed chamber. 

Length 0.7 mm., breadth 0.25 mm. Cat. No. 324625a, 6, c, U.S.N.M. 

Specimens of this species occurred in the Gatim formation at the 
following three stations, U.S.G.S. No. 6033c, marl from second bed 
from bottom, just below lower clay, Gatun Section, relocated line 
Panama Railroad; U.S.G.S. No. 6035, in gray-green, fine-grained, 
sandy shell marl vicinity of Mindi Hill, and U.S.G.S. No. 6036, in 
dark-colored fine-grained, sandy clay marl, at Monkey Hill, Mount 
Hope Station. 

At none of these stations were more than a few specimens found 
but all seem referable to this species. 

Family LAGENIDAE. 
Genus LAGENA Walker and Boys, 1784. 

LAGBNA STRIATA (d'Orbiffur), rur. 8TRUM0SA R«aM. 

Plate 21, fi?. 7. 

Lagena strumosa Reuss, Zeitschr, geol. Ges., 1858, p. 434; Sitz, Akad. 

Wlss. Wlen, vol. 46, pt. 1, 1862 (1863), p. 328, pi. 4, fig. 49. 
Lagena striata (d'Obbigny), var. strumosa Cushman, Bull. 71, U. S. Nat. 

Mus., pt. 3, 1913, p. 20, pi. 7, figs. 7-10. 

Description. — ^Test clavate or subglobular, the body portion orna- 
mented with numerous longitudinal raised costae, apical end with a 
single stout spine; neck short and stout, typically with a phialine lip 
and transverse costae. 

Diameter 0.5 mm. Cat No. 324626, U.S.N.M. 

A single specimen of this variety was obtained in material from 
U.S.G.S. No. 6010, from the lower part of the Gatun formation, dark 
clay, north of Pedro Miguel Locks. This is the only representative 
of the genus in the whole series of samples examined. The speci- 
men lacks the neck except the base and the tip of the apical spine. 



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OBOLOOY AND PALEONTOLOGY OF THE CANAL ZONE. 59 

GeniiB NODOSARIA Lamarck, 1812. 

NODOSARIA COMMUNIS d'Orbigny. 

Plate 21, fig. 8. 

Nodosaria {Dentalina) communis d'Obbigny, Ann. Sci. Nat., vol. 7, 1826, 

p. 254, No. 85. 
Kodosaria commurUs Riruss, Verst. B<5hm. Kreld., pt. 1, 1845. p. 28, pi. 12, 

fig. 21.— H. B. Beady, Rep. Voy. Challenger^ Zoologj% vol. 9, 1884, p. 504, 

pi. 62, figs. 19-22. 

Description. — ^Test elongated, slender, gradually tapering, slightly 
curved, chambers slightly inflated in the middle, sutures distinct, 
slightly depressed, somewhat oblique ; wall smooth. 

Length 2 mm. ? Cat No. 324627. 

A single fragment showing four chambers was obtained in material 
from U.S.G.S. No. 6036, Oatun formation, from dark-colored fine- 
grained, sandy clay marl at Monkey Hill, Mount Hope Station. The 
fragment with its general characters, its smooth surface, slightly 
inflated chambers and oblique sutures seem to clearly indicate this 
species. 

NODOSABIA INSECTA Schwager? 

Plate 21, n^. 9. 

Nodosaria insecta Schwageb, Norara Exped. Geol. Thiel., pt. 2, 1866, p. 224, 
pi. 5. figs. 52, 53. 

Description. — Test elongate, gradually tapering from the nearly 
acute slender base to a broad apical end, which is the greatest in 
diameter of any of the chambers of the test; chambers numerous, 
inflated, nearly spherical, sutures much depressed; wall smooth, 
apertures with a slight neck and circular opening. 

Length 2.3 mm. Cat. No. 32462&Z, 6, U.S.N.M. 

Specimens were found in the lower part of Culebra formation both 
at U.S.G.S. No. 6010, in dark clay, north of Pedro Miguel Locks, 
and 6012a, from lower dark clay beneath lower conglomerate, one- 
fourth .mile south of Empire Bridge. 

. The specimens are very close to the species described by Schwager 
from the Tertiary of Kar Nicobar. The two forms, megalospheric 
and microspheric. occur in the Panamanian material, the latt«r being 
much more slender at the initial end than in the megalospheric. 

NODOBABIA BAPHANI8TBUM (LtauuMW). 

Plate 21, fig. 10. 

XautUuM raphaniatrum Linnaeus, Syst Nat, ed. 10, 1758, p. 710. 
Xodonnria raplwnistrvm Revss, in Oelnltz. Grundr. Versteiu, 1845-46, p. 

653, pi. 24, Apr. 6. — Jones, Parker, and H. B. Brady, Monogr. Pal. Soc., 

vol. 19. 1866, p. 50, pi. 1, figs. 6-8. 
8370*'— 18— Bull. 103 6 



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60 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Description. — ^Test elongate, subcylindrical, slightly tapering, 
chambers numerous, distinct, apertural end with a short tapering 
neck; wall ornamented with longitudinal costae continued clear to 
the aperture, about 12-15 in number. 

Length 4 mm. Cat. No. 324629, U.S.N.M. 

A single specimen of this species figured here was obtained fronr 
U.S.G.S. No. 6010, lower part of the Culebra formation, in dai4: 
clay, north of Pedro Miguel Locks. The specimen is not complete 
at the initial end but the last six chambers including the aperture 
are very well preserved. 



NODOfl 

Plate 21, ^g, 11. 

A fragment consisting of one complete chamber and the adjacent 
parts of two others was found in the same material, U.S.G.S. No. 
6010, as the above but nearly twice the diameter. The costae are 
also more numerous. Without further material it is imsafe to try 
to determine the fragment, but the occurrence of another species at 
this station should be at least recorded. Cat. No. 624630, n.S.N.&£ 

Geniu CRISTELLARIA Lamarck^ 1812. 

CBISTBLLABIA BOTULATA (Lamarck). 
Plate 22, fig. 1. 

''Ck>mu Hammonis seu NautlU" Plancub, Ck>nch. Mln., 1789, p. 18, pi. X 

fig. III. 
Lentimlites rotulata Lamarck, Ann. Mus., vol. 5, 1804, p. 188, No. 8; vol. 8, 

180C. pi. 62, fig. 11. 
CHsteUaria rotulata d'Orbiony, Mem. Soc. 6^1. France, ser. 1. vol. 4, 1840, 

p. 26. pi. 2. figs. 16-18.— H. B. Bbady, Rep. Voy. Challenger, Zoology, 

vol. 9, 1884, p. 547, pi. 69, figs. 13a, 6. 

Description, — Test comparatively large, biconvex, close coiled 
throughout, chambers variable in number in the coil, sutures distinct, 
periphery not lobulated, usually not keeled; previous apertures of 
the test usually visible as is often the preceding coil at least in part; 
wall smooth. 

Diameter up to 2 mm. Cat. Nos. 324631a, 6, (?, d^ «, U.S.N.M. 

This seems to be the commonest species in the Panamanian materiaL 
It differs slightly in form in the various stations but all may be 
grouped under this species. It occurred in two groups of stations 
as noted in the chart of distribution. They are as follows: Lower 
part of the Culebra formation at U.S.G.S. No. 6010, in dark 'clay, 
north of Pedro Miguel Locks; No. 6012a, in lower dark clay beneath 
lower conglomerate, one-fourth mile south of Empire Bridge. 
Gatun formation at U.S.G.S. No. 6033c, in marl from second bed 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 61 

from bottom just below lower clay, Gatun section, relocated line 
Panama Bailroad; No. 6035, in gray green, fine-grained, sandy shell 
marl, vicinity of Hindi Hill; and No. 6036 in dark-colored, fine- 
grained, sandy clay marl of Monkey Hill, Mount Hope Station. 

CRISTELLABIA ITALICA (Defnmce). 

Saracenaria itafica Defranck, Diet Scl. Nat, vol. 32, 1824, p. 177; vol. 47, 

1827, p. 344; Atlas Conch., p!. 13, flg. 6. 
Cristellaria (Saracenaria) italica d'Orbigny, Aun. Sci. Nat, vol. 7, 1826, 

p. 298, No. 26 ; Modules, Nos. 19 and 85. 
Cristellaria italica Parkeb, Jones, and H. B. Br.\dy, Ann. Mag. Nat Hist, 

ser. 3, vol. 10, 1865, pp. 21, 32, pi. 1, figs. 41, 42.— H. B. Bhadt, Rep. 

Voy. ChMcnger, Zoology, vol. 9, 1884, p. 544, pi. 68, figs. 17, 18, 20-23. 

Description. — ^Test with the early portion close coiled, later por- 
tion more or less uncoiled, chambers numerous, those of the last- 
formed portion being triangular in cross section, periphery keeled, 
and the apertural face broad and flattened, the sides angled and ex- 
tending on either side to the keel in flat faces, sutures but slightly 
depressed, wall smooth ; apertures peripheral, radiate, usually with no 
nedc 

Diameter 0.75 mm. Cat. No. 324632, U.S.N.M. 

Two specimens are evidently of this species in a young stage^ the 
uncoiling not yet having proceeded to a great degree. They are from 
TJ.S.G.S. No. 6086, Gatun formation, in dark-colored, fine-grained, 
sandy clay marl from Monkey Hill, Mount Hope Station. 

CRISTELLARIA PROTUBERANS, new ipMiM. 

Plate 22. flg. 2. 

Description. — Test compressed, close coiled, biconvex, seven cham- 
bers in each coil, each much inflated in its central portion, space be- 
tween much compressed, flattened, periphery sharply and broadly 
keeled; aperture peripheral, radiate. 

Diameter 0.80-1.20 mm. 

Three specimens of this species occurred at TJ.S.G.S. No. 6010, 
lower part of Culebra formation, in dark clay north of Pedro Miguel 
Locks. It is in some respects similar to species found in the Western 
Pacific, especially in comparatively deep water off the Philippines. 

Type-specimen.— G2Lt No. 324633, U.S.N.M. 

CRISTELLARIA VAUGHANI. new spedM. 

Plate 22, flg. 3. 

Description, — Test much compressed, with a slight tendency to un- 
coiling in the last-formed chambers, periphery slightly keeled, not 
lobulated, rounded, about nine chambers in the last-formed whorL 



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62 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

sutures slightly curved backward, extending in to the umbilicus so 
that only the last-formed coil is visible from the exterior, surface 
smooth except for linos of beads along the sutures extending from 
the umbilicus to the periphery; apertural face truncated or even 
slightly concave, aperture radiate, peripheral, with a short cylin- 
drical neck. 

Diameter 0.75 mm. 
The type-sections of this species are from U.S.G.S. No. 6035, 
Gatun formation, in gray green, fine-grained, sandy shell marl from 
the vicinity of Mindi Hill. It also occurred at 6036, Gatun forma- 
tion, in dark-colored, fine-grained, sandy clay marl from Monkey 
Hill, Mount Hope Station; No. 6019/, upper part of Culebra forma- 
tion, fourth limy bed from bottom, section opposite Las Cascadas, 
Gaillard Cut; and No. 6010, lower part of Culebra formation, in 
dark clay, north of Pedro Miguel Locks. 

This species is somewhat sug.aestive of some forms < f C\ wethereliij 
but has no longitudinal ribbing. It is perhaps nearest to C. geminata 
described by Brady from the Philippines and South Sea Islands, but 
lacks the typical papillate surface common in that species. 

The species is named for Dr. T. Wayland Vaughan, whose collec- 
tions in the Canal Zone have added much to the available forairdni- 
fera from this region. 

Type-specvmem.—CKt. No. 324634, U.S.N31. 

Genus UYIGERINA d'Orbifirny, 1826. 

UVIGBRINA CAKARIENSIS d'Orbiffny. 

Plate 21>, fig:. 5. 

" Testae plnelforme minuBCulae " Soldani. Testaceographla, vol. 2, 1798» 

p. 18, pi. 4, figs. E, F, G, H. 
Uvigerina nodosa, var. B d*Orbigny, Ann. Scl. Nat., vol. 7, 1826, p. 209, No. 3. 
Vrigerina canarienaia d'Orbigny, Foram. Canaries, 1889, p. 138, pi. 1, flgs. 

2r)-27.~H. B. Brady, Rep. Voy. Challenger, Zoology, vol. 9. 1884, p. 573. 

pi. 74, figs. 1-3. 

Description. — Test elongate, chambers numerous, spirally arranged,' 
triserial, inflated, separated by distinct sutures; wall smooth except 
for the early chambers which may show traces of spines or longitudi- 
nal striae; apertural end usually with a tubular neck and often a 
phialine lip. 

Length 0.75 mm., diameter 0.35 mm. Cat. No. 324635, U.S.N.M. 

The only typical material of this species is from U.S.G.S. No. 6035, 
Gatun formation, in gray-green, fine-grained sandy shell marl from 
thft vic»initv of Mindi Hill. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 63 

UVIGESINA CANARIBNSI8 d'OrMgnsr. Tariety. 

Plate 22, fig. 6. 

A larger and much stouter, entirely smooth variety as shown in the 
above figure was found in material from U.S.G.S. No. 6010, lower 
part of Culebra formation, in dark clay, north of Pedro Miguel 
I^ks. Cat. No. 324636, U.S.N.M. 

UVIGERINA PYGMAEA d'OrUsny. 

Plate 22, fig. 4. 

" Polymorpha Pinelforniia " Soi:dant, Testaceographla. vol. 1, pt. 2, 1791, 

pi. 130. tigs. SH, tt. 
I'vigcrina plgmea d'Orbiony, Auu. Sei. Nat., vol. 7, 1826, p. 260, pi. 12, figs. 

8, 9; Modules, 1826, No. 67. 
Uiigcrina pygmaca d'Okbigny, Foram. Foss. Bass. Tert. Vienne, 1846, p. 190, 

pi. 11. figs. 2o, 26.— H. B. Bbady, Rep. Voy. Challenger, Zoology, vol. 9, 

1884, p. 575, pi. 74, figs. 11-14. 

Description. — Test subcylindrical, triserially spiral, chambers nu- 
merous, inflated, sutures deep; wall ornamented by numerous longi- 
tudinal costae, those of each chamber usually independent of those 
of adjacent chambers; aperture with a short cylindrical neck and 
phialine lip. 

Length 0.75 mm., breadth 0.32 mm. Cat. No. 324637a, 6, c, 
U.S.N.M. 

Specimens referable to this species occurred in the Culebra forma- 
tion at U.S.G.S. No. 6012^ in lower dark clay beneath lower con- 
glomerate, one-fourth mile south of Empire Bridge, Gaillard Cut, 
and No. 6012e? in clay and sandstone just below conglomerate at 
base of green clay one-half to three- fourths of a mile north of Con- 
tractors Hill, Gaillard Cut. 

Specimens of a slightly different character were abundant at No. 
^5, Gatun formation, in gray-green, fine-grained sandy shell marl, 
vicinity of Mindi Hill. 

UVIGERINA TBNUISTRIATA RevM. 

Plate 22, fig. 7. 

Vvigerina striata Reuss, SUz. Kals. Akad. Wiss. Wlen, vol. 52, 1870, p. 485.— 
VON ScHLicHT, Foram. Pletzpuhl, 1870, pi. 22, figs. 34r-86. — ^H. B. Bbady, 
Rep. Voy. Challenger, Zoologj% vol. 9. 1884, p. 574, pL 74, figs. 4-7. 

Description. — ^Test subcylindrical, chambers spirally arranged, tri- 
serial at least in the early portion, later portion sometimes biserial 
and more slender; chambers inflated, sutures deep, walls ornamented 
by numerous longitudinal costae, except the last chambers, which tend 
to become smooth or nearly so ; aperture with a short tubular neck 
and often a phialine lip. 



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64 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Length 0.85 mm., breadth 0.30 mm. Cat. No. 324638, U.S.N.M. 

Specimens referred to this species were very common in material 
from U.S.G.S. No. 6036, Gatun formation, in dark-colored, fine- 
grained sandy clay marl, from Monkey Hill, Mount Hope Station. 
Many of the specimens become almost uniserial in the last-formed 
portion. 

Genus SIPHOGENERINA Schlnmberger, 188S. 

SIPHOGENERINA RAPHANUS (P«rk«r and Jones) Tmr. TRANSVERSU8, new TartotF. 

Plate 22, fig. S, 

Description. — ^Test subcylindrical, compased of comparatively few 
chambers, the earlier ones spirally arranged, later and greater por- 
tion of the test uniserial, sutures very prominently indented, be- 
tween the longitudinal costae, aperture with a short cylindrical neck. 

Length, 1.25 mm. ; diameter, 0.54 mm. Cat. No. 324646, U.S.N.M. 

This variety differs from the typical form in the much greater 
prominence of the transverse depressions marking the sutures, occa- 
sionally as in the figure suggesting the depressions of S. dimoryhcu 
The specimens were frequent in material from U.S.G.S. No. 6010, 
lower part of the Culebra formation, in dark clay, north of Pedro 
Miguel Locks. 

Fanuly GLOBIGERINIDAE. 
Genus 6L0BI6ERINA d'Orbifirny, 1826. 

GLOBIGERINA BULLOmES d'OrUgny. 

GloJHgerina hulloidea d*Obbiony, Ann. Sci. Nat, vol. 7, 1826, p. 277, No. 1 ; 
Modules, No. 17, and No. 76; in Barker, Webb, and Berthelot. Hist. 
Nat Isles Canaries, 1839, pt 2, Foraniinif6res, p. 132, pi. 2, figs. 1-3, 
28.— H. B. Beady, Rep. Voy. Challenger, Zoology, vol. 9, 1884, p. 59S, 
pi. 77 ; pi. 79, figs. 8-7. 

Description. — Test subglobose, spiral, visible portion composed of 
but few chambers from below, usually three to five, all visible from 
the dorsal side, sutures deep, chambers inflated, umbilicate below; 
surface reticulate ; aperture single, from each chamber, of good size 
opening into the central umbilical cavity on the ventral side. 

Diameter, 0.60 mm. Cat Nos. 324639-45. 

Specimens referable to this widely distributed species were ob- 
tained from the following stations: In the Culebra formation, 
U.S.G.S. No. 6009, from black clays and sandy beds at lower end 
of Pedro Miguel Locks; 6010 in dark clay, north of Pedro Miguel 
Locks; 6019/, in fourth limy bed from bottom, Las Cascadas sec- 
tion, Gaillard Cut In the Gatun formation, U.S.G.S. No. 6029&, 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 65 

in argillaceous and sandy indurated marl, one-fourth to one-half 
mile north of Camp Cotton on relocated line, Panama Railroad; 
6033c in marl from second bed from bottom, just below lower clay, 
Gatun Section relocated line, Panama Railroad; 6035, in gray green, 
fine grained, sandy shell marl, near Mindi Hill ; and 6036, in dark 
colored, fine grained, sandy clay marl, Monkey Hill, Mount Hope 
Station. 

The specimens from the last three stations are very well preserved 
and in fact might almost be recent material, while those of the other 
stations were fragmentary, often glauconitic. G. bulloides^ var. 
triloba Reuss was occasional in the last three stations where the 
genus was really very conmion. 

GLOBIGERmA INFLATA d'OrMgnr. 

Qiobigerina inflata d'Orbiqny, in Barker, Webb, and Berthelot, Hist. Nat. 
Isles Canaries, vol. 2, pt 2, 1839, Foraminiftees, p. 134, pi. 2, figs. 
T-O.—EL B. Brady, Rep. Voy, ChaUenger, Zoology, vol. 9, 1884, p. 601, 
pi. 79, figs. 8-10. 

Description. — ^Test composed of numerous inflated chambers usu- 
ally arranged in a spiral test with about three volutions, the last- 
formed one with four chambers, dorsal side of test nearly flat, ventral 
side extended, especially in the last-formed whorl ; ventrally umbili- 
cate; surface finely reticulate; aperture large, opening toward the 
umbilicus. 

Diameter, 0.75 mm. Cat. Nos. 324647, 8, 9, U.S.N.M. 

Specimens occurred at U.S.G.S. No. 6010, lower part of the Cule- 
bra formation, in dark clay north of Pedro Miguel Locks; and in 
the Gratun formation at the last two of the stations already referred 
to, namely, 6035 and 6036. 

GLOBIGEBINA DUBIA Egger. 

Olobigerina dubia Egger, Neiies Jtihrb. flir Min., 1857, p. 281, pi. 9, flffs. 
7-9.— H. B. Bbady, Rep. Voy. Challenger, Zoolojo', vol. 9, 1884. p. 595, 
• pi. 79. figs. 170-0. 

Description. — ^Test composed of numerous inflated chambers ar- 
ranged in a nauilloid spiral all visible from above, the last coil only, 
consisting of 5 to 6 chambers, visible from below, ventral side with a 
central umbilicus, surface reticulate; apertures opening into the 
central umbilical cavity. 

Diameter 0.75 mm. Cat. Nos. 324650-54. 

At the following stations specimens referable to this species were 
found: Culebra formation, U.S.G.S. No. 6010, in dark clay, north 
«f Pedro Miguel Locks; 6025, in dark, hard, sandy clay about 200 



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66 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

yards south of southern end of switch at Bohio Bidge Station, re- 
located line, Panama Railroad. Gratun formation, U.S.G.S. No. 
6033<?, in marl from second bed from bottom, just below lower clay, 
Gatun Section, relocated line, Panama Bailroad; 6035, in gray green, 
fine grained, sandy shell marl near Mindi Hill and 6036 in dark 
colored, fine grained, sandy clay marl, Monkey Hill, Mount Hope 
Station. 

As in the case of the preceding species the specimens from the 
last three stations were very finely preserved while those of the 
others were glauconitic 

GLOBIGEBINA CONGLOBATA H. B. Btadjr. 

OloMgerina conglobata H. B. Brady, Quart. Journ. Mlcr. Sci., vol. 19, 1879, 
p. 72; Rep. Voy. Challengei^ Zoologj-, vol. 9, 1884, p. 603, pi. 80, figs. 
1-5; pi. 82, fig. 5. 

Description, — Test subglobular, early chambers arranged in a com- 
pact spiral, the last three chambers in the complete adult test form- 
ing nearly the whole of the visible portion of the test, wall coarsely 
reticulat'e ; main aperture at the inner margin of the chamber with 
several rounded secondary apertures along the margins of the 
chamber where it is attached to adjacent ones. 

Diameter up to 1 mm. Cat. Nos. 324655-6. 

Specimens of G. conglohata were found in small numbers in the 
Gatun formation at stations 6035 and 6036. They were typical but 
perhaps hardly as well developed as in some Recent material. Its 
occurrence here is rather interesting as it is almost unknown in the 
fossil condition. 

GLOBIGERINA SACCULIFERA H. B. Brady. 

t 
GJohigerina helicina Carpenter (not G. helicina d'Orblgny). Intr. Foram., 

1862. pi. 12, fig. 11. 

Olohigerina sacculifera H. B. Brady, Geol. Mag., Dec. 2, voL 4, 1877, p. 

535 ; Quart. Journ. Micr. Sci., vol. 19, 1879, p. 73 ; Rep. Voy. Challenger, 

Zoology, vol. 9, 1884, p. 604, pi. 80, figs. 11-17 ; pi. 82, fig. 4. 

Description. — Test composed of nmnerous chambers, in its early 
stages very similar to G. huUoides but later developing a more oblong 
form, the chambers extended, somewhat compressed and with ac- 
cessory apertural openings, the final chamber often flattened and 
irregularly formed toward the outer end; wall strongly reticulat^ed 
in all but the final chamber which is much smoother than the others; 
aperture large, arched, with other accessory openings in the chambers 
of adult specimens. 

Diameter up to 1 mm. Cat. Nos. 324657-8, U.S.N.M. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 67 

Specimens were not uncommon in material from the Gatun forma- 
tion at stations 6035 and 6036. As in the case of G. conglohata the 
specimens were hardly as well developed as they are in recent speci- 
mens, but nevertheless had the characteristic marks of the speciea 
As in G. conglohata the records of this species are almost entirely 
limited to Recent material, its occurrence as a fossil being practically 
unknown. 

GLOBIGBRINA AEQUILATERAUS H. B. Bndr. 

CasHduUna globuJosa (part) Egger, Nouos Jahrb. filr Mln., 1857, p. 296, 

pi. 11, fig. 4. 
Olobifferina aequilateralis H. B. Brady, Quart. Journ. Micr. Scl., voL 19, 

1879, p. 71 ; Rep. Voy. Challefiper, Zoology, vol. 9, 1884, p. 605, pi. 80, 

figs. 18-21. 

Description. — ^Test composed of numerous inflated chambers, ar- 
ranged in a planospiral manner, at least the last formed coil, cham- 
bers increasing rapidly in size as added, usually 5 to 6 in the last 
formed volution; sutures depressed, periphery lobulated; surface 
reticulate; aperture large, at the base of the inner margin of the 
<?hamber. 

Diameter up to 1 mm. Cat. Nos. 324659-61, U.S.N.M. 

In the material from the Gatun formation at three stations, Nos. 
6033^, 6035, 6036, this species was not uncommon. The only char- 
acter in which there seems to be a difference from the Recent material 
is in the early chambers which occasionally show at one side as a flat 
spiral while the later chambers are bilateral. The species is not a 
oonmion one as a fossil. 

GenuB ORBULINA d'OrUgny, 1839. 

ORBULINA UNTVERSA d'OrMgny. 

OrbiUina univrrsa d'Obbigny, In De la Sagra, Hist. Fls. Pol. Nat Cuba, 
1839, ForaminlWres, p. 3, pi. 1, fig. 1.— H. B. Brady, Rep. Voy. Chal- 
lenger, Zoology, vol. 9, 1884, p. 6aS. pi. 78; pi. 81, figs. 8-26; pi. 82, 
figs. 1-3. 

Description. — ^Test in adult form typically consisting of a single, 
spherical visible chamber, which may or may not have contained 
within the early Globigerine stages ; wall strongly reticulate, a single 
large circular aperture and smaller openings at the base of each 
reticulaticm. 

Diameter up to 1 mm. Cat. Nos. 324662-8, U.8.N.M. 

Specimens were not uncommon in the Gatim formation at the 
three stations, Nos. 6083c, 6085, and 6036. Occasional specimens 
show the double form as figured by Brady. The specimens other- 
wise are like the common run of Becent material. 



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68 BULLETIN 103^ TJNITBD STATES NATIONAL MTJSBXTM. 

Family ROTAUIDAE. 
Genus DISCORBIS Lamarck, 1804. 

DI8CORBI8 OBTUSA (d'OrMffnr). 

Plate 23, Aks. la^. 

Ronalina obtusa d'Orbiont, Foram. Foss. Bass. Tert Vieime, 1846, p. 179, 

pi. 11, flgs. 4-6. 
DUoorbina obtusa H. B. BsAinr, Rep. Voj. ChaUenger, Zoology, voL 9, 1884, 

p. 644, pi. 91, flgs. 9o-c. 
Discorbis obtusa Gushman, Bull. 71, U. S. Nat. Mus., pt. 5, 1915, p. IS, 

flgs. 12a-c. 

Desoinption. — Test biconvex, dorsal side more so than the ventral 
side, peripheral margin roimded ; chambers comparatively few, about 
five in the last formed whorl; sutures curved, depressed; wall per- 
forate; aperture an elongate narrow slit extending from the um- 
bilicus nearly to the periphery. 

Diameter 0.60 mm. Cat. No. 324664, U.S.N.M. 

The only station from which this species was obtained is U.S.G.S- 
No. 6850, from Pleistocene marl near Mount Hope, a quarter mile 
from the present sea beach and 6 to 8 feet above high tide. 

Genus TRUNCATUUNA d'OrUgny, 1826. 

TRUNCATUUNA AMERICANA, new ipeeicii. 

Plate 23, flgs. 2a~v. 

Description. — Test nearly plano-convex; ventral side strongly 
convex, periphery keeled, dorsal side nearly flat; chambers numerous, 
up to nine in the last formed coil ; sutures curved, prominent, slightly 
limbate, umbilicate below; surface smooth, aperture nearly pe- 
ripheral. 

Diameter 0.65 mm. 

Type-specimen. — (Cat. No. 324665, U.S.N.M.) from the upper 
part of the Culebra formation, at U.S.G.S. No. 6019/, fourth limy 
bed from bottom. Las Cascades se<;tion, Gaillard Cut. 

TRUNCATUUNA PTGMEA Hantkra. 

Plate 28, flgs. 8a^. 

Trunoatulina pygmea Hantken, Mitth. Jahrb. ung. geoL Abstalt., vol 4, 

1875, p. 78, pi. 10. flg. a 
TrunoatuUna pygmaea H. B. Bbadt, Rep. Voy. OhdUenger, Zoology, vol. 9^ 

1884, p. 666, pi. 95, flgs. 9, 10. 



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QBOLOQY AND PALEONTOLOGY OF THE CANAL ZONE. 69 

Description. — Test nearly equally biconvex, peripheral margin 
bluntly rounded; chambers numerous, the sutures oblique, distinct, 
often limbate ; aperture a narrow slit extending from near the periph- 
ery nearly to the umbilicus. 

Diameter 0.65 mm. Cat. No. 324666-7, U.S.N.M. 

The only station at which this species occurred is in the upper part 
of the Culebra formation, U.S.G.S. No. 6019^, upper part of second 
hard, limy, sandstone bed. Las Cascadas section, Gaillard Cut. It 
is rather larger than the usual i-un of T. pygmaea but is evidently 
this species. 

Specimens from the Gatun formation, U.S.G.S. No. 6036, while 
having fewer chambers and somewhat larger size are questionably 
referred here. One specimen is figured on plate 24, figure 2. 

TRUNCATUUNA UNGERIANA (iTOrbicny). 
Plate 24, fig. 1. 

R^UUina ungeriana d'Obbigny, Foram. Foss. Bass. Tert. Vienne, 1846, p 

157, pi. 8, figs. 16-18. 
Planorbnlina ungeriana H. B. Br^vdy, Trans. Liun. Soc. London, vol. 24, 

1864, p. 469, pi. 48, fig. 12. 
TnmcatvHnu ungeriana Reuss, Denkschr. Akad. Wiss. Wien, vol. 25, 1865, 

p. 161.— H. B. Beady, Rep. Voy. Challenger, Zoology, vol. 9, 1884, p. 664. 

pi. 94, figa 9ar-d. 

Description.— Test biconvex, dorsal side less convex than the ven- 
tral; peripheral margin subacute, slightly carinate, chambers nimier- 
ous, 10 to 12 in the last formed whorl, sutures distinct, slightly lim- 
bate on the dorsal side; aperture a narrow arched opening running 
ventrally from the peripheral margin. 

Diameter 0.50 mm. Cat. Nos. 324668-9, U.S.NJ^f. 

Specimens referable to this species but not entirely typical were 
obtained in material from lower part of the Culebra formation, as 
follows: U.S.G.S. No. 6009, from black clays and sandy beds at lower 
end of Pedro Miguel Locks; and 6012a, from lower dark clay be- 
neath lower conglomerate, one-fourth- mile south of Empire Bridge, 
west side Gaillard Cut, below Culebra. 

TRUNCATUUNA WUELLERSTORFI (Sehwager). 
Plate 24, fig. 3. 

AnomaUna umellerstar/l Schwageb, Novara Exiwd., geol. Thell.. vol. 2, 

1866, p. 258, pi. 7, figs. 105, 107. 
TruncaUaina icuellerstorfl EL B. Bbadt, Rep. Voy. Challenger, Zoology, vol. 

9, 1S84, p. 662, pL 93, figs. 8, 9. 

Description. — ^Test plano-convex, dorsal side nearly flat, ventral 
side slightly convex; chambers numerous, elongate, curved; sutures 



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70 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

strongly curved, somewhat limbate, periphery bluntly rounded, 
slightly lobulated, especially near the apertural end of the IsLst 
formed coil; wall coarsely punctate; aperture peripheral, a short 
curved opening. 

Diameter of larger specimens slightly more than 1 nun. 

Numerous very typical specimens of this species occurred in mate- 
rial from the lower part of the Culebra formation, U.S.G.S. No. 
<)010, from dark clay, north of Pedro Miguel Locks. Less typical 
specimens occurred in the upper part of the Culebra formation at 
U.S.G.S. ^012d, from clay and sandstone just below conglomerate 
at base of green clay, west side of Gaillard Cut, below Culebra ; and 
6019/, from fourth limy bed from bottom, Las Cascadas section, 
Gaillard Cut. 

Cat. Nos. 324670-2, U.S.N.M. 

TRUNCATULINA CULEBRENSIS, new ipeciM. 
Plate 24, figs. 4a, 5. 

Description. — ^Test biconvex, much compressed, peripheral margin 
rounded ; chambers nimierous, as many as thirteen in the last formed 
coil, long and narrow, gently curved, sutures broad, limbate, smooth. 
the areas between very coarsely punctate; apertural face of chamber 
somewhat depressed, flattened, the carinate borders extending out 
beyond at either side; aperture a narrow slit situated at the base of 
the chamber on the periphery. 

Diameter up to 1.6 mm. 

The only occurrence of this species was in the tipper part of the 
Culebra formation, U.S.G.S. No. 6012^, from top part of limy sand- 
stone below upper conglomerate near foot of stairs, west side Gail- 
lard Cut 

This, a large and striking species, in some of its characters sug- 
gesting T. wuelleratorfi but, as will be seen by a comparison of the 
figures of the two, really very different. 

Type-specimen^— Cdit. No. 324673, U.S.N.M. 

Genus PULVINULINA Parker and Jones, 1862. 

PULVINUUNA 8AGRA (d'Orbiffny) . 

Plate 24, figs. 6a, b. 

Rotalina sagra d'Orbignt, In De la Sagra, Hist Fis. Pol. Nat Ouba, 1839. 
Foraminif^res, p. 77, pi. 5, figs. 18-15. 

Description, — ^Test ovate, biconvex, the vwitral side more convex 
than the dorsal, peripheral margin subacute, carinate; chambers com- 
paratively few in number increasing rapidly in size in the last formed 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 7 1 

ones, the last formed chamber on the ventral side making up a large 
part of the area of the test, sutures distinct, curved, sli^tly de- 
pressed, more so on the ventral side; wall smooth except for the usual 
fine punctations; aperture ventral near the umbilicus. 

Length 0.60 mm., breadth 0.40 mm. Cat No. 324674. 

The only record for this species from Panama is from the Gatun 
fonnation, U.S.G.S. No. 6035, in gray green, fine grained, sandy 
?hell marl, near Mindi Hill. This species, described by d'Orbigny 
from Cuba, seems to be a common species in the American Miocene. 

PULVINUUNA CONCBKTRICA Parker and Jones. 

Plate 25. fig. 1. 

PulvUtiUina concentrica (Parker and Jones, MS.) H. B. Brady, Trans. Linn. 
Soc. London, vol. 24, 1864, p. 470. pi. 48, fig. 14.— H. B. Brady, Rep. 
Voy. Cliallenger, Zoology, vol. 9, 1884, p. 686, pi. 105, figs. lo-c. 

Description, — Test biconvex, oval; peripheral margin rounded; 
chambers comparatively few, usually seven in the last formed coil, 
sutures covered by clear shell material joining with the carinal border 
and often covering a large portion of the test, both above and below, 
especially toward the center; wall smooth, finely punctate; aperture 
a narrow slit on the peripheral portion of the ventral side. 

Diameter 1.2 mm. Cat. No. 324675, U.S.N3I. 

The only specimen of this species is from the Gatun formation, 
U.S.G.S. No. 6035, in gray green, fine grained, sandy shell marl near 
Mindi Hill. The specimen as will be seen from the figure is very 
typical. 

PULVINULINA lUENARDU (d'Oibigny). 

Plate 2.'>, figs. 2, 3. 

Rotalia tnenardU d'Obbiqny, Ann. Sci. Nat, vol. 7, 1826, p. 273, No. 26; 

Modules, No. 10. 
Pulvinulina menardii Owen, Journ. Linn. Soc. London (Zool.), vol. 9, 1867, 

p. 148, pi. 5, fig. 6. — H. B. Brady, Rep. Voy. Challenger, Zoolojry. vol. 9, 

1884, p. 690, pi. 103, figs. 1, 2. 

Description. — ^Test plano-convex, ventral side convex, doreal side 
nearly flat; compressed, umbilicatc; peripheral margin thin, slightly 
lobulated, carinate; chambers five or six in the last formed coil; 
sutures distinct, limbate and broad on the dorsal side, curved, on the 
ventral side more depressed, not limbate, nearly straight; wall 
smooth, finely punctate; a])erturo extending peripherally from the 
umbilicus, usually with an overhanging lip. 

Diameter up to 1 mm. Cat. Nos. 324676-8, U.S.N.M. 

Specimens apparently belonging to this species so widely dis- 
tributed in the present oceans were obtained in the Gatun formation 
at U.S.G.S. No. 6035 in gray green, fine grained, sandy shell marl, 



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72 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

vicinity of Mindi HiH ; and 6036 in dark colored, fine grained, sandy 
clay marl from Monkey Hill, near Mount Hope Station. A figure of 
one of these is shown in plate 25, figure 3. From 6033^, Gatun 
formation, in marl from second bed from bottom, just below lower 
clay, Gatim section, relocated line of the Panama Railroad, are even 
more typical specimens, one of which is here figured on plate 25, 
figure 2. 

Genus SIPHONINA Reuss, 1849. 

SIPHONINA RETICULATA (Csjaek). 

Plate 24, flg. 5. 

Rotalina reticulata Czjzsk, Haldinger*s Nat Abh., vol. 2, 1848, p. 145, 

pi. 13, figs. 7-©. 
Siplumina reticulata Bbowit, Lethaea C^gnostica, ed. 3, vol. 3, 1853^66, 

p. 227, pi. 35 (?), figs. 23(i-c.— CusHMAN, Bull. 71, U. S. Nat Mus., 

pt 5, 1915, p. 43, flg. 48; pi. 16, flg. 4; pi. 28. flg. 3. 
TruneatuUna reticulata H. B. Brady, Rep. Voy. Challenger, Zoolog>% vol. 9, 

1884, p. 669, pi. 96, figs. 5-8. 

Description. — ^Test biconvex, ventral side slightly more so than 
the dorsal, peripheral margin acute, carinate; chambers numerous 
rather indistinct, sutures slightly depressed, curved; wall rather 
coarsely perforate; aperture peripheral with a short, broad neck and 
somewhat flaring phialine lip. 

Diameter 0.65 mm. Cat. No. 324679, U.S.N.M. 

The only station at which this species occurred is in the Gatun 
formation, U.S.G.S. No. 6036, in dark colored, fine grained, sandy 
clay marl of Monkey Hill, Mount Hope Station. 

Although the specimen is not perfectly preserved the tubuli of the 
peripheral margin are lacking as is the case in some large recent 
specimens. 

Family NUMMULTnDAE. 

GeniM NONIONINA d'Orbignj, 1826. 

NONIONINA DBPRESSULA (WallMr and Jacob). 
Plate 25, figs. 5a, 6. 

Nautilus depresaulus Walkeb and Jacob, Adam's Essays, Kanmacher's 

ed., 1798, p. 641. pi. 14, flg. 83. 
Nonionina depressula Parkeb and Jones. Ann. Mag. Nat. Hist., ser. 8, vol. 4, 

1859, pp. 339, 841.— H. B. Brady, Rep. Voy. Challenger, Zoology, vol 

9, 1884, p. 725, pi. 109, figs. 6, 7.— Bagq, Bull. U. S. Geol. Surv., No. 513, 

1912, p. 88. pi. 26, figs. 16<i-c; pi. 28, figs. 7, 8. 

Description. — ^Test more or less rounded in side view, slightly 
elongate, about ten chambers in the last formed coil, apertural view 
narrow, periphery broadly rounded, sides nearly parallel, about two 
and a half times as high as broad, umbilicus slightly depressed, 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 78 

usually filled with secondary shell material and a slight extension 
peripherally along the sutures which are slightly depressed; aper- 
ture a narrow curved slit. 

Diameter 0.60 mm. Cat Nos. 824680-1, U.8.N.M. 

Distribution. — Specimens of this species occurred in the Gatun 
fonnation at U.S.G.S. No. 6033c, in marl from second bed from 
bottom, just below lower clay, Oatun Section, relocated line of the 
Panama Bailroad ; and 6035, in gray green, fine grained, sandy shell 
marl, vicinity of Mindi Hill. The specimens are rather typical, per- 
haps varying in the direction of increased length from most recent 
specimens. 

NONIONINA 8GAPHA (FltdMl and MoU). 

Plate 25, figs. 6a, b. 

NautUits Bcapha Fichtel and Moll, Test. Micr., 1808, p. 105, pi. 19, figs. d-/. 

Ifonitmina Bcapha Pabkeb and Jones, Ann. Mag. Nat. Hist, ser. 3, vol. 5, 
1860, p. 102, No. 4.— H. B. Braot, Nat. Hist. Trans. Northumberland 
and Durham, vol. 1, 1865, p. 106, pi. 12, figs. 10a, 6.— H. B. Brady, Rep. 
Voy. ChaUenger, Zoology, vol. 9, 1884, p. 730, pi. 109, figs. 14. 15, and 
16. — ^H. B. Bbady, Pabkeb, and Jones, Trans. Zool. Soc., vol. 12, 1888, p. 
280, pi. 43, fig. 20.— WooDWABD and Thomas. Geol. Nat. Hist Surv. 
Minnesota, vol. 3. 1893, p. 48, pi. E, figs. 35, 36.— Eogeb, Abh. k5n. Bay. 
Akad. Wiss. Mttnchen. CI. II. vol. 18, 1893, p. 424, pi. 19, figs. 43, 44.— 
Gofis, Kongl. Svensk. Vet. Akad. Handl., vol. 25, 1894, p. 104, pi. 17, 
fig. 830.— MoBTON, Proc. Portland Soc. Nat. Hist. vol. 2. 1897. p. 121, 
pi. 1. fig. 23.--FUKT, Ann. Rep. U. S. Nat. Mus., 1897 (1899), p. 337. 
pi. 80, fis- 1. — Fobnasini, Mem. Accad. Sci. 1st Bologna, ser. 6, vol. 1, 
1904, p. 12. pi. 3, fig. 4 ; pi. 13, fig. 5.— Millett. Journ. Roy. Mlcr. Soc.. 
1904. p. 601.— Baog. Proc. U. S. Nat. Mus.. vol. 34, 1908. p. 164.— Side- 
BOTFOM, Mem. and Proc. Manchester Lit. and Philos. Soc.. vol. 53, No. 
21. 1909. p. 13 ; vol. 54. No. 16. 1910, p. 29, pi. 3, fig. 13.— Bagg, Bull. 
U. S. Geol. Sun-. No. 513, 1912. p. 88, pi. 27, figs. 1-8. 
Polystomella crispa, var. {Nonionina) acapha Pabkeb and Jones, Philos. 
Trans., vol. 15.5, 1865, p. 404. pi. 14, figs. 37. 38 ; pi. 17, figs. 55, 56. 

Description. — ^Test in side view longer than wide, about ten cham- 
bers in the last formed coil, rapidly increasing in length as added, 
sutures evenly curved, slightly depressed, periphery broadly rounded, 
in apertural view the face of the last formed chamber making up a 
large part of the visible surface, wall smooth, finely pimctate, some- 
what umbilicate ; aperture an arched slit at the base of the chamber. 

Length 0.60 mm. Cat. No. 324682, U.S.N.M. 

Specimens of this species were collected in the Gatun formation 
at a single station, U.S.G.S. No. 6033t?, in marl from second bed 
from bottom, just below lower clay, Gatun section, relocated line of 
the Panama Railroad. 

The specimen figured in apertural view was placed to show the 
aperture rather than the full size of the apertural face which is really 



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74 BULLETIN 103^ UNITED STATES NATIONAL MUSEUM. 

larger than appears in this view, the earlier portion of the coil being^ 
narrow. 

NONIONINA PANAMBNSIS. im^ 



Plate 26, figs, la, 5. 

Description. — ^Test in side view subcircular, last formed chamber 
composed of about nine chambers, in front view bilaterally sym* 
metrical, rapidly increasing in breadth as chambers are added, aper- 
tural face of chamber broadly rounded, early portion slightly keeled ; 
sutures rather strongly curved, slightly limbate, slightly depressed ; 
wall smooth, distinctly punctate ; aperture a narrow curved slit at the 
base of the apertural face of the chamber. 

Diameter 0.65 mm. 

Specimens of this species were obtained from the lower part of 
the Culebra formation, U.S.G.S. No. 6010, north of Pedro Miguel 
IxK^s, in dark clay. 

Type-specimen.— Cvii. No. 324683, U.S.N.M. 

NONIONINA ANOMALINA, mw tpedM. 

Plate 26, figs. 2a. 6. 

Description, — Test in side view nearly circular, deeply umbilicate* 
peripheral margin broadly rounded, bilaterally symmetrical, about 
seven chambers in the last formed coil, sutures little if at all de- 
pressed, indistinct, last formed chambers extending but part way 
across the test, tending toward alternating arrangement; aperture 
a narrow slit at the base of the chamber. 

Diameter 1.26 mm. 

Type-specimen. — (Cat. No. 324684, U.S.N.M.) from the lower part 
of the Culebra formation, in dark clay, north of Pedro Miguel Locks 
(U.S.aS. No. 6010). 

The last two chambers suggest CassidvUna^ but the similarity does 
not continue further. 

Genmi POLTSTOMELLA Lamarck, 1822. 

POLTSTOMBLLA 8TRIATO-PUNCTATA (FltdMl and Moll). 

Plate 26, flgB. 8a, 5; 4a, h. 

NautUus striato-punctatus Fichtel and Moll, Test Mlcr., 1808, p. 61, pi. 9,. 

figs. (l^. 
Polystomella striato-punctata Parker and Joines, Ann. Mag. Nat Hist, ser. 

3, vol. 5, 1860. p. 103, No. 6.— H. B. Brady. Rep. Voy. ChaUmger^ 

Zoology, vol. 9, 1884, p. T33, pi. 109, figs. 22, 23. 

Description. — ^Test bilaterally qrmmetrical, subcircular in side view^ 
umbilicate, peripheral margin broadly rounded, eight to ten chambers 



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GBOLOGY AND PALBONTOLOQY OF THE OANAL ZONE. 75 

in the last formed coil; sutures slightly curved, depressed; wall 
smooth, distinctly punctate; septal lines with regularly arranged, 
narrow bridging; aperture a narrow semicircular opening at the base 
of the apertural face of the chamber, showing occasionally traces of 
division into a series of smaller openings. 

Diameter 0.50 to 0.65 mm. Cat Nos, 324685-7, U.S.N.M. 

Specimens were obtained in the Culebra formation, U.S.G.S. No. 
6020a, opposite Las Cascadas, in lowest fossiliferous bed, third bed 
below lowest limestone. These were very largely glauconitic, and 
of the form figured in 4a, ft. The species was also found in the 
Gatun formation, U.S.G.S. No. 6029a, one- fourth to one-half mile 
north of Camp Cotton, relocated line of the Panania Railroad, in 
Uie softer sandy marls at the base of the section. The form figured 
in 3a, J, is from a Pleistocene deposit at U.S.G.S. No. 5850, loose 
shells and marl from near Mount Hope, one-fourth mile from present 
beach, 6 to 8 feet above high tide. 

POLTSTOMELLA SAGRA d'Orblsny. 

Plate 26, figs. 5a, h. 

Polyatomella Sagra d*Orbigny, In De la Sagra, Hist. Fis. Pol. Nat. Cuba, 
1839, Foraminifdres, p. 55, pL 6, figs. 19, 20. 

Description. — ^Test bilaterally symmetrical, subcircular in side 
new; peripheral margin rounded, ten or more chambers in the last 
formed coil; sutures distinct, curved, slightly depressed in the last 
fonned portion, not at all depressed in the early part of the coil; 
early half of the coil with definite raised, longitudinal ribs, corre- 
sponding to the bridging over the sutures, persisting longest on the 
peripheral portion of the test, later portion smooth; bridging of 
earliest portion of coil regular, short, in the last formed sutures in- 
creasing considerably in length; apertural face smooth, punctate; 
roughly triangular in outline, the angles rounded; aperture a very 
narrow slit at the base of the apertural face of the chamber. 

Diameter 0.65 mm. Cat. No. 324688, U.S.N.M. 

The only station at which this species was obtained is a Pleistocene 
deposit at U.S.G.S. No. 5860, loose shells and marl from near Mount 
Hope, one-fourth mile from present beach and 6 to 8 feet above high 
tide. 

A comparison of this figure with the original given by d'Orbigny 
in his Lao., .-monograph will show the very striking similarity be- 
tween the Cuba and Panama specimens, and I have no hesitation in 
referring this material to d'Orbigny's species. 
8370'--18— Bull. 103 6 



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76 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

POLTBTOMBLLA MACBLLA (FIchtol tMi IMl). 

Plate 27, figs, la, &. 

Nautilus maoellu8t var. a, Fiohtel and Moll, Test. Micr., 1803, p. 66, pi. 10, 

figs. e-ij. 
Polyatotnella macella Pabkeb and Jokes, Ann. Mag. Nat Hist, ser. 3, vol. 5, 

1860, p. 104, No. 8. — ^H. B. Bbady, Rep. Voy. Challenifer, Zoology, vol. 9, 

1884. p. 737. pL.110. figs. 8, 9, 11. 

Description. — Test compressed, bilaterally symmetrical, peripheral 
margin acute, somewhat carinate, not lobulated, sixteen to twenty 
chambers in thp last formed coil; reticulated bridgings occupying a 
greater area than the intermediate portions; umbilical region slightly 
depressed, with a few large pores; aperture a curved or V-shaped 
slit at the base of the apertural face, either simple or divided into 
secondary openings. 

Diameter, 0.75 mm. Cat. Nos. 324689-90, U.S.N.M. 

Specimens were obtained, from two stations in the Emperador 
limestone, as follows: U.S.G.S. 6016, from cream-colored coral lime- 
stone, old quarry one-quarter mile north of west from Empire ; and 
6016, one-third mile north of west of the same place. 

POLTSTOMELLA CRISPA (Linnseiu). 
Plate 27, figs. 2a, h. 

"Cornu Hammoiiis orblculatnm " Plancus, Conch. Min., 1739, p. 10, pi. 1, 

fig. 2. 
Nautilus crispus Linnaeus, Syst Nat, ed. 12, 1767, p. 1162. 
Polystomella crispa Lamarck, Anlm. sans Vert, vol. 7, 1822, p. 625, No. 1. — 

d'Orbigny, Foram. Foss. Bass. Tert Vlenne, 1846, p. 125, pi. 6, figs. 9- 

14.— H. B. Brady, Rep. Voy. Challenger, Zoology, vol. 9, 1884, p. 736, 

pi. 110, figs. 6, 7. 

Description, — Test bilaterally symmetrical, much compressed, pe- 
ripheral margin obtusely angled; umbilical region not depressed; 
chambers numerous, eighteen to twenty chambers in the last formed 
coil, sutures indistinct, bridging wider than the intermediate clear 
space; margin not lobulated; umbilical region imibonate, filled with 
clear shell material, often with a few pores; aperture a narrow slit 
at the base of the apertural face of the chamber, usually showing 
more or less division into secondary openings. 

Diameter, up to 1.25 mm. Cat. No. 324691, U.S.N.M. 

Specimens referable to this species were obtained from the Gatun 
formation at U.S.G.S. No. 60296, one-fourth to one-half mile north 
of Camp Cotton on relocated line of the Panama Railroad, indurated 
argillaceous and sandy marl. 



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GEOLOGY AKD PALEONTOLOGY OP THE CANAL ZONE. 77 

POLTSTOMELLA CRATICULATA (Ftehtd and IMl). 

Plate 27, figs. 3o, &. 

Ntiutilus craticulatus Fichtel and Moll, Test. Micr., 1803, p. 51, pL 5, figs. 

h-k. 
PolystomeUa oratioulata d'Orbigny, Ann. Sci. Nat., voL 7, 1826, p. 284, 

No. 3.— Cabpenteb, Intr. Foram., 1862, p. 279, pL 16, figs. 1, 2.--H. B. 

Bbady, Rep. Voy. Challenger, Zoology, vol. 9, 1884, p. 739, pi. 110, figs. 

16, 17. 

Description^ — ^Test bilaterally symmetrical, somewhat compressed; 
peripheral margin broadly rounded; not lobulated, chambers very 
numerous, forty or more in the last formed coil, narrow; umbilical 
region filled with clear shell material with numerous pores; bridged 
area about equal to that between; aperture a series of openings at 
the base of the apertural face. 

Diameter, 1 mm. Cat. No. 324692, U.S.N.M. 

This species was found in considerable numbers in the Culebra 
formation at U.S.G.S. No. 6025, in foraminiferal marl and coarse 
sandstone about 200 yards south of the southern end of the switch at 
Bohio Kidge station, relocated line, Panama Railroad. 

The specimens have not as subglobose a form as many recent 
specimens, but in other respects the characters are very similar. 

POLTSTOMELLA, species? 

Numerous stations have a species of Polystomella which is very 
much like P. sagra and yet is not so definitely characterized as are 
the si>ecimens of that species from station 6025. 

The stations at which this form of Polystomella occurs are in the 
lower part of the Culebra formation at U.S.G.S. No. 6009, black 
clays, six or seven hundred feet south of Miraflores Locks. In Las 
Cascadas section, Gaillard Cut, 6019 J, from the 4 feet of dark strati- 
fied tuflF and clay overlying the lower limestone bed; 6019/, from 
fourth limy bed from bottom; 6020a, from the lowest fossiliferous 
bed. Li the Emperador limestone at 6015 and 6016 from cream- 
colored coral limestone, old quarry, one-quarter mile north of west 
from Empire. In the Gatun formation at 6029a, from lowest hori- 
zon, one- fourth to one-half mile north of Camp Cotton. 

Cat Nos. 824693-8,'U.S.N.M. 

Genus AMPHISTEGINA d'OrUgny, 1826. 

AMPmSTEGINA LESSONn <rOrbiffiiy. 

Amphistcgina lessonii d'Okbigny, Ann. Sci. Nat., vol. 7, 1826, p. 304, No. 8, 
pi. 17, figs. 1-4 ; Modules, No. 98.— H. B. Bbady, Rep. Voy.. Challenger, 
Zoology, vol. 9, 1884, p. 740. pi. Ill, figs. 1-7. 

Description. — ^Test lenticular, usually more convex on one side than 
the other ; composed of about twenty-five chambers in the last formed 



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78 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

coil, wall smooth except near the aperture on the ventral side where 
there is usually a papillose area of greater or less extent; periphery 
usually somewhat rounded; sutures on the dorsal side with a single 
simple angle ; below usually divided into two deep lobes by deep con- 
strictiona 

Diameter, 1-2.5 mm. Cat. Nos. 324699-08, U.S.N.M. 

This species is common in the lower horizons of the area occurring 
at the following stations: Culebra formation, 6009, 6012a, rf, 6019<?, d^ 
6027; Emperador limestone, 6015, 6016; Gratun formation, 6029a, 6, c. 

At some of these stations specimens are rather frequent. In the 
matrix this species may often be indistinguishable in a superficial ex- 
amination from worn centers of Orbitoids or Nummulites. It is a 
common Tertiary species. 

Family MIUOLIDAE. 
Genus QUINQUELOCUUNA d'Orbigny, 1826. 

QUINQUELOCULINA SEMINULUM (Unnmeiu). 

Plate 27, figs. 4a, 6; plate 28; plate 29, figs. la-e. 

Serpula seminulum Linnaeus, Syst. Nat., ed. 10, 1758, p. 786; ed. IS 

(Gmelin). 1758, pp. 37, 39. 
QuinquelocuUna seminulum d'Obbigny, Ann. Scl. Nat., vol. 7. 1826, p. 30B, 

No. 44. 
MUiolina seminulum Williamson, Rec. Foram. Great Britain, 1858, p. 85, 

pi. 7, figs. 183-189.— -H. B. Brady, Rep. Voy. Challenger, Zoology, vol. 9, 

1884, p. 157, pi. 5, figs. 6a, &, c. 

Description, — ^Test oval in front view; thickest in the middle, 
visible exterior composed of five chambers, three visible from one 
side and four from the other, sutures slightly depressed, distinct; 
wall smooth, periphery rounded, aperture somewhat contracted, 
usually with a single simple tooth. 

Length about 1 mm. Cat. Nos. 324709-13, U.S.N.M. 

Very typical specimens were obtained from U.S.G.S. No. 5850, 
among loose shells and marl from near Mount Hope, from ditch 
through swampy ground, one- fourth mile from present sea beach and 
6 to 8 feet above high tide (Pleistocene). Specimens very similar 
but slightly more rotund were obtained from the Gatun formation. 
No. 6036, in dark colored, fine grained, sandy clay marl, at Monkey 
Hill, Mount Hope Station. 

Varietal forms here figured and which may be referred to Q. semi- 
nulwra were obtained from the Culebra formation at No. 6010, from 
dark clay, north of Pedro Miguel Locks; 6019a, a single specimen 
from lower limestone of Las Cascadas section; 6025, a single glau- 
conitic specimen from foraminiferal marl about 200 yards south of 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 79 

southern end of switch at Bohio Eidge Station, relocated line of the 
Panama Sailroad. Another much flattened glauconitic specimen 
from this last station is also referred here. 

QUINQUELOCULINA CONTOBTA <rOiibigii7. 

Plate 29, figs. 2a-c. 

Quinqueloculina contorta d*Oebiqny, Foram. Fobs. Bass. Tert, Vienne, 1846, 
p. 298, pi. 20, figs. 4-6. 

Description. — Test about twice as long as broad, chambers rather 
narrow and elongate, in end view polygonal, peripheral margin 
broadly curved, sides nearly at right angles to the peripheral face 
with a sharp angle at the junction ; sutures deep, apical end and initial 
end of final chamber truncated ; aperture rounded with a single tooth ; 
wall smooth. 

Length 0.65 mm. Cat. No. 324714, U.S.N.M. 

The only material of this species was obtained from U.S.G.S. 5850, 
among loose shells and marl, from near Mount Hope, from ditch 
through swampy ground, about one-fourth mile from present sea 
beach and 6 to 8 feet above high tide (Pleistocene). 

QUINQUELOCULINA AUBERIANA d'Orbiipiy. 

Plate 29, figs. 3a-c. 

Qumqueloculina auheriaiia d'Obbignt, in De la Sagra, Hist. Fis. Pol. Nat. 

Cuba, 1839, ForaminlfSres, p. 193, pi. 12, figs. 1-3. 
Milidlina auberiwia H. B. Bradt, Rep. Voy. Challenger, Zoology, vol. 9, 1884, 

p. 162, pL 5, figs. 8, 9. 

Description. — Test sli^tly longer than broad, periphery of the 
diambers angled with a concave area at each side of the angle, sutures 
somewhat depressed, distinct; wall smooth; aperture with a single, 
usually simple, occasionally slightly bifid tooth. 

Length about 1 mm. Cat. No. 324715, U.S.N.M. 

Two specimens of this species were obtained in material from 
U.S.G.S. 6860, among loose shells and marl, from near Mount Hope^ 
from ditch through swampy ground, about one-fourth mile from pres- 
ent sea beach and 6 to 8 feet above high tide (Pleistocene) . This is a 
common species of the shallow- water littoral of tropical seas. 

QUINQUELOCULINA UNDOSA Karrer. 

Plate 30, figs. la-e. 

Quinqueloculina undo$a Kabrer, Sitz. Akad. Wiss. Wlen, vol. 58, abth. 1, 

1868, p. 150, pi. 3, fig. 1. 
MUiolhui undoaa Kabbkb, Rep. Voy. Challenger^ Zoology, vol. 9, 1884, p. 176, 
pi. 6, figs. 6-8. f 

Description^ — ^Test elongate, two or two and a half times as long as 
wide; chambers sub-polygonal, the angles more or less irregular giv- 



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80 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

ing an undulate appearance to the chambers, apertural end typically 
with a slightly projecting neck, aperture with a single tooth; wall 
smooth. 

Length 1.25 mm. Cat Nos. 324716-17, U.S.N.M. 

Specimens referable to this species were obtained in the Emperador 
limestone, at U.S.G.S. 6016, from old quarry, one-third mile north 
of west of Empire; and in the Culebra formation, at 6025, in 
foraminiferal marl about 200 yards south of the southern end of 
the switch at Bohio Ridge Station, relocated line, Panama Bailroad. 

The specimens are not so contorted as in some recent ones but show 
characteristic undulations of the chamber borders. 

QUINQUBLOCULINA BICORNIS (Walker and Jacob). 

Plate 30, figs. 2a-c; 3a, b. 

"Serpula blcornis ventricosa," Walkkb and Boys, Test Mln., 1784, p. 1, 

pi. 1. fig. 2. 
" Frmnentaria foeniculum*' Soldani, Testaceographia, vol. 1, pt 3, 1T96, 

p. 229, pi. 154, figs. &&, cc. 
SerpuUb bicomiB Walker and Jacob, Adams's Essays, Kanmacher** ed., 

1798, p. 633, pi. 14, fig. 2. 
MilioUna bicomis Williamson, Rec. Foram. Great Britain, 1858, p. 87, pi. 

7, figs. 190-192.— H. B. Brady, Rep. Voy. Challenger, Zoology, vol. 9, 

1884, p. 171, pi. 6, figs. 9, 11, 12. 

Description. — ^Test in side view about twice as long as wide; 
futures rather deep, distinct, chambers more or less keeled, wall 
ornamented with numerous rather fine longitudinal raised costae; 
aperture slightly exserted, rounded, with a single tooth. 

Length 0.75 mm. Cat. Nos. 324718-9, U.S.N.M. 

Specimens were obtained in a Pleistocene deposit at U.S.G.S. 5850, 
among loose shells and marl, from near Mount Hope, from ditch 
through swampy ground about one-fourth mile from present sea 
beach, and 6 to 8 feet above high tide. 

From the Culebra formation, U.S.G.S. 6025, in foraminiferous 
marl about 200 yards south of southern end of switch at Bohio Ridge 
Station, relocated line, Panama Railroad, were obtained, rather 
poorly preserved and somewhat glauconitic specimens, but showing 
traces of a longitudinal series of raised ridges. They are question- 
ably referred here and one is figured, on plate 30, figure 3. 

QUINQUELOCULINA PANABIENSIS, new spedet. 

Plate 31, figs. la-c. 

Description. — ^Test nearly as wide as long, the last formed chamber 
tending to become loose coiled, growing &,way from the preceding 
ones on the apertural half of the inner margin, apertural end free. 



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GEOLOGY AlH) PALEONTOLOGY OF THE CANAL ZONE. 81 

peripheral margin broadly rounded, sutures much depressed; wall 
smooth; aperture circular. 

Length 0.85 mm. 

This species was obtained from the Gatun formation, U.S.G.S. 
6036, in dark colored, fine grained, sandy clay marl, from Monkey 
Hill, Mount Hoi)e Station. 

It is imusual in the breaking away of the last formed chamber 
from the original close coiled method of growth. 

Type-8peoimen.—CB,t. No. 324720, U.S.N.M. 

Genus SIGMOILINA Schlumberger, 1887. 

SIGMOIUKA TENUIS (Cijsek). 

Plate 31, figs. 4ct-c. 

Quinqueloculina tenuis Ozjzek, Haidlnger*s Nat. Abhandl., yol. 2, 1847, p. 

149, pi. 13, figs. 31-«4. 
Spiroloculina tenuis H. B. Bbady, Rep. Voy. Challenger, Zoology, Yol. 9, 

1884, p. 152, pi. 10, figs. 7-11. 
SigmoUina tenuis Sidebottom, Mem. and Proc. Manchester Lit and Phllos. 

Soc., vol. 48. No. 5, 1904, p. 6. 

Description. — ^Test about twice as long as wide, narrow, compressed, 
visible chambers 5 or 6 on either side, chambers, narrow, roimded, 
sutures depressed, distinct; wall smooth, aperture exserted, rounded. 

Length 0.66 mnL Cat. Nos. 324721-8, U.S.N.M. 

Specimens of this species were obtained in the Gatun formation 
at the following three stations : U.S.G.S. 6033t?, in marl from second 
bed from bottom, just below lower clay, Gatun section, relocated line 
of the Panama Kailroad ; 6035, in gray green, fine grained, sandy 
diell marl, vicinity of Mindi Hill; and 6036, in dark colored, fine 
grained, sandy clay marl, from Monkey Hill, Mount Hope Station. 
, These three stations have several species in common as will be 
seen by a glance at the accompanying chart of distribution. 

SIGMOIUKA A8PEBULA (Kamr). 
Plate 31, figs. 3 a, &. 

Sptroludna asperula Kabbeb, Sitz. Akad. Wiss. Wien, yol. 57, 1868, p. 186, 
pi. 1, fig. 10.— H. B. Brady, Rep. Vol. Challenger, Zoology, vol. 9, 1884, 
p. 152, pi. 8. figs. 13, 14, and 11. 

Description. — ^Test but slightly longer than wide, very much com- 
pressed, sutures somewhat indistinct, several chambers visible from 
each of the flattened sides; wall covered with fine arenaceous parti- 
cles; aperture exserted, nearly circular. 

Length, 0.8 mm. Cat. Nos. 324724-5, U.S.N.M. 



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82 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Specimens were not uncommon in the Gatun formation at two 
stations, U.S.G.S. 6035, in gray green, fine grained, sandy shell marl, 
vicinity of Mindi HiU, and 6036, in dark colored, fine grained, sandy 
clay marl, from Monkey Hill, Momxt Hope Station. 

Genus TRILOCUUNA d'OrUgny, 1826. 

TRILOCULINA TRIGONULA (Lammrck). 

Plate 32, fijj. 1. 

MUiolites trigonula Lamarck, Ann. Mus., vol. 5. 1804, p. 351, No. 3. 
TrUoculina trigonula d'Orbigny, Aim. Sci. Nat., vol. 7, 1826, p. 299, No. 1. 

pi. 16, figs. 5-9. 
MUiolina trigonula Williamson, Rec. Porara. Great Britain, 1858. p. 88, 

pi. 7, figs. 180-182.— H. B. Br^^dy, Rep. Voy. Challenger, Zoology, vol. 9, 

1884, p. 164, pi. 3, figs. 14-16. 

Description. — Test in apertural view triangular, angles rounded, 
chambers rapidly increasing in size as added, but three visible in 
adult test; outer wall broadly rounded, in front view oval, sutures 
distinct, aperture not produced, lip and tooth indistinct. 

Length, 0.75 mm. Cat. No. 324726, U.S.N.M. 

A single specimen referable to this species occurred at U.S.G.S. 
5850, in Pleistocene marly material from near Mount Hope, one- 
fourth mile from present sea beach and about 6 to 8 feet above high 
tide. 

This is a common species in shallow water of recent oceans. 

TRILOCULINA TRICARINATA d'OrblKny. 

Plate 32. fi?. 2. 

TrilocuHna tricarinata D'ORmoNY, Ann. Scl. Nat,, vol. 7, 1826, p. 209. No. 7 ; 

Modules, No. 94. — H. B. Brady, Trans. lArxn, Soc. London, vol. 24, 1864, 

p. 446. pi. 48, fig. 3. 
MUiolina tricarinata H. B. Brady, Rep. Voy. Challenger, Zoology, vol. 9; 

1884, p. 165, pi. 3, figs. 17a, &. 

Description. — Test differing from T. trigomda largely in the an- 
gles, which are acute, the sides concave, at least toward the borders, 
center of the side either flat or slightly convex, in end view rather 
sharply triangular, in front view oval ; neck slightly produced, aper- 
ture rounded, tooth wanting in this specimen. 

Length, 0.60-0.70 mm. 

Four specimens were collected in the Culebra formation at U.S.G.S. 
No. 6025, foraminiferal marl about 200 yards south of southern end 
of switch at Bohio Bidge Station, relocated line, Panama Railroad. 

Two of the four specimens had the neck somewhat elongated, the 
others were more nearly normal in this respect. The specimens were 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 83 

somewhat altered and showed traces of apparently a glau(jonitic 
interior. 

TRILOCULINA BULBOSA, new ipcdM. 

Plate 32, fig. 3. 

Description. — ^Test from exterior composed of three visible cham- 
hers. the last formed one making the largest part of the test. The 
next to the last about half the size of the last and the first formed 
one very small in comparison, test in end view nearly biloculine, with 
the last formed chamber nearly as wide as the whole test in its great- 
est width, in front view breadth and height about equal, chambers 
very rotund, sutures deep, aperture without a neck, rounded, tooth 
indistinct or wanting. 

Length, about 0.65 mm. 

Type-specimen.— {CvX. No. 324728, U.S,N.M.) from the Gatun 
formation, U.S.G.S. Station 6029a, lowest horizon, one- fourth to one- 
half mile north of Camp Cotton on relocated line Panama Railroad. 
Another specimen was obtained, also in the Gatun formation, at No. 
6030, from fossiliferous marl, from cut on north side of swamp 1^ 
miles north of Monte Lino, relocated line of the Panama Railroad. 

In each case a single somewhat glauconitic specimen was obtained. 
The species has the last two chambers developed greatly, the third 
one very small, the whole test appearing almost biloculine. The 
specimens from the two stations were practically identical. 

TRILOCUUKA PROJECTA, new ipedet. 

Plate 33, fig. 1. 

Description. — Test in end view composed of three radially pro- 
jecting portions, the intervening portions deeply concave, in side 
view about as long as wide, sutures somewhat indistinct, periphery 
broadly rounded; wall covered with a thick encrustation of sand 
grains giving the whole exterior a decidedly arenaceous appearance; 
aperture with a slightly projecting neck and phialine lip ; apertural 
opening circular, in the specimen figured without a distinct tooth. 

Length 0.75 mm. 

Type-specimen. — (Cat. No. 324729, U.S.N.M.) From gray green, 
fine grained, sandy shell marl from vicinity of Mindi Hill, U.S.G.S. 
No. 6035, Gatun formation. 

This is an interesting modification of this genus, comparable in 
the structure of the test to Quinqueloculina agglutinans d'Orbigny 
and others of the same character. 



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84 BULLETIN 103, UNITBD STATES NATIONAL MUSEUM. 

Genus BILOCULINA d'OrUgny, 1826. 

BILOCULINA BULLOIDES d'OrUcny. 

Plate 88, fig. 2. 

BUocuUna huttoides d'Orbigny, Ann. Scl. Nat, vol. 7. 1826, p. 297, No. 1, 
pi. 16, figs. 1-4; Modules, No. 90. — H. B. Bbadt, Rep. Voy. Challenger, 
Zoology, Tol. 9, 1884, p. 142, pi. 2, figs. 5, 6. 

Description. — ^Test with but two visible chambers in the adult, in 
end view, each semicircular, in front view elliptical, very rotund, 
inflated, suture distinct; aperture usually nearly circular, some^vhat 
produced. 

Length 0.60 mm. Cat. No. 324730, U.S.N.M. 

The only specimen of this species is from the Gatun formation, 
U.S.Q.S. 6036, from dark colored, fine grained, sandy clay marl from 
Monkey Hill, Mount Hope Station. • 

Genus SPIROLOCUUNA d'OrUgny, 1826. 

SPIROLOCULINA EXCAVATA cTOrblgny. 

Plate 81, fig. 2. 

Spiroloculina excavata d'Obbignt, Foram. Foss. Bass. Tert Vlenne, 1846, 
p. 271, pi. 16, figs. 19-21.— H. B. Brady, Rep. Voy. Challenger, Zoology, 
vol. 9, 1884, p. 161, pi. 9, figs. 5, 6. 

Description. — ^Test planospiral, chambers much elongated, thickest 
at the basal end, apertural end slightly produced, central portions 
much excavated, due to the gradual increase in the width of the 
chambers as added; periphery somewhat convex, angles rounded; 
sutures distinct except toward the center; apertural end produced, 
aperture rounded; tooth wanting in our specimen. 

Length 1.4 mm. Cat. No. 324731, U.S.N.M. 

Specimens were obtained at Station No. 5850, in marl of Pleisto- 
cene, Mount Hope, Canal Zone, by D. F. MacDonald. 

Genofl ORBICULINA Lamarck, 1816. 

ORBICUUNA ADUNCA (Flditel and Moll). 

Plate 88, fig. 8. 

Nautilus orbiculus Fichtel and Moll, Test. Mlcr., 1803, p. 112, pi. 21. 
Orbiculina adunca Lamarck, Tabl. Encycl. et M^tli., 1816, pi. 468. flga 

2a~c.— H. B. IJrady, Hop. Voy. Challenger, Zoology, vol. 9, 1884, p. 209, 

pi. 14, figs. 1-13. 

Description. — ^Test planospiral, chambers very long, divided into 
simple chamberlets, sides with alar projections extending nearly to 
the umbilicus, sutures distinct; apertures numerous, peripheral. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 85 

Diameter 1.6 mm. Cat. No. 324782, U.S.N.M. 

A few specimens were obtained from U.S.G.S. 5850, from Pleisto- 
cene marl near Mount Hope, about one-fourth mile from present 
sea beach and about 6 to 8 feet above high tide. 

EXPLANATION OF PLATES. 

Plate 19. 

ri6. 1. Tewtviaria ahbreviata d'Orblgny. X 50. a, apertural view; 6, front 
view. 

2. TextuUiria soffittula Defrance. X 80. a, apertural view ; b, front view. 

3. Temtularia agglutinans d'Orblgny. X 50. a, apertural view; &, front 

view. 

4. Textularia laminata, new species. X 80. a, apertural view; b, front 

view. 

5. Textularia suhagghitinana, new species. X 85. a, apertural view; &, 

front view, 
d. Textularia carinata d'Orbigny. X 50. a, apertural view ; 6, front view. 

Plate 20. 

Fig. 1. Textularia panameniis, new species. X 65. a, ai)erture view ; I), front 
view. 

2. Chrysalidina pulchella, new species. X 110. a, apertural view; &, 

\iewed from flat side ; c, viewed from angle. 

3. Oaudryina triangularis Oushman. X 85. a, apertiural view; &, front 

view. 

4. Oaudryina flintii Gushman. X 50. 

5. Clavulina parisiensis d*Orbigny. X 85. 

6. Clavulina communis d'Orblgny. X 85. 

Plate 2L 

Fig. 1. Bolivkia, species. X 65. 

2. Bolivina aenariensis (Ck)sta). X 65. 
3. Bolivina cf. B. punctata d*Orblgny. X 65. 

4. Bolivina robusta H. B. Brady. X 185. 

5. Bigenerina nodosaria d*Orbigny. X 27. a, apertural view; b, front 

view. 

6. Virgulina squamosa d*0rblgny. X 65. 

7. Lagena striata (d'Orbigny), var. strumosa Reuss. X 65. 

8. Nodosaria communis d'Orbigny. X 65. 

9. Nodosaria cf. A", insecta Schwager. X 35. 

10. Nodosaria raphanAstrum (Linnaeus). X 85. 

11. Nodosaria, species ? X 85. 

Plate 22. 

Fig. 1. Crist ellaria rotulata (Lamarck). X 85. 

2. Cristellaria protuberans, new species. X 65. 

3. Cristellaria vaugJiani, new species. X 65. 

4. Uvigerina pygmaea d'Orblgny. X 65. 

5. Uvigerina canariensis d*Orbigny. X 65. 

6. Uvigerina canariensis d'Orbigny var. X 65. 

7. Uvigerina tenuistriata Reuss. X 65. 

8. Sipliogenerina raphanus (Parker and Jones) var. transverstts. new 

variety. X 85. 



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86 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Plate 28. 

FiQ. 1. Discorbis ohtuaa (d'Orblgny). a, dorsal view; &, ventral view; c, pe- 
ripheral view. X 65. 

2. Truncatulina americana, new species, a, dorsal view; &, ventral view; 

c, peripheral view. X 65. 

3. TruncatuUna pygmea Hantken. a, dorsal view; &, ventral view; c, pe- 

ripheral view. X 65. 

Plate 24. 

Fig. 1. Truncatulina unperiana (d'Orbigny.) X 65. 

2. Truncatulina cf. T. pygmea Hantken. X 33. 

3. Truncatulina wuellerstorfi (Sch wager). X 50. 

4. Truncatulina culehrensis, new species. X 33. a, dorsal view; 6, pe- 

ripheral view. 

5. Siptumina reticulata (Gzjzek). X 65. 

6. Pulvinulina sagra (d'Orbigny). X 65. a, dorsal view; &, ventral view. 

Plate 25. 

Fig. 1. Pulvinulina concentrica Parker and Jones. X 85. 
2. Pulvinulina menardii (d*Orbigny). X 65. 
a Pulvinulina menardU (d'Orbigny). X 65. 

4. PuMnuUna, species ? X 66. 

5. Nonionina depres8ula (Walker and Jacob). X 65. a, side view; b, 

apertural view. 

6. Nonionina acapha (Fichtel and Moll). X 65. o, «ide view; &, aper- 

tural view. 

Plate 26. 

Fig. 1. Nonionina panamensis, new species. X 65. a, side view ; b. apertural 

view. 
2. Nonionina anomalina, new species. X 65. a, side view; b, apertural 

view. 
8. Poly&tomeUa striato-punctata (Fichtel and Moll). X 65. a. side view; 

6, apertural view. 

4. Polystomella atriato-punctata (Fichtel and Moll). X 65. a, side view; 

6, apertural view. 

5. Poly$tomella sagra d*Orbigny. X 65. a, side view; ft, apertural view. 

Plate 27. 

Fig. 1. Polystomella macella (Fichtel and Moll). X 65. a, side view; 6, aper 
tural view. 

2. Polystomella crispa (Linnaeus). X 35. a, side view; 6, apertural view. 

8. Polystomella craticulata (Fichtel and Moll). X 50. a, side view; 6, 
apertural view. 

4. Quinqueloculina seminulum (Linnaeus). X 65. a, rear view; &, aper- 
tural view. 



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GEOLOGY AISTD PALEONTOLOGY OF THE CANAL ZONE. 87 

Plate 28. 

Fi&l. Quinqueloculina semin/ulum (Linnaeus). X 65. a, front view; &, rear 
view ; o, apertaral view. 

2. Quinqueloculina seminulum (Linnaeus). X 180. a, front view; &, rear 

view; c, apertnral view. 

3. Quinqueloculina aeminulum (Linnaeus) var. X 05. a, front view; b, 

rear view; c, apertural view. 

Plate 29. 

FI6.L Quinqueloculina aeminulum (Linnaeus) var. X 80. a, front view; b, 

rear view; c, apertural view. 
' 2. Quinqueloculina contorta d'Orbigny. X 65. a, front view ; b, rear view ; 
c, apertural view. 
3. Quinqueloculina auberiana d*Orbigny. X 65. a, front view; b, rear 
view; c, apertural view. 

Plate 80. 

Fia 1. Quinqueloculina undosa Karrer. X 50. a, front view ; &, rear view ; o, 
apertural view. 
2. Quinqueloculina bicomis (Walker and Jacob). X 65. a, front view; b, 

rear view ; c, apertural view. 
3w Quinqueloculina bicomis (Walker and Jacob)?. X 50. a, front view; 
5, rear view. 

Plate 81. 

Fi6. 1 Quinqueloculina panamenHSf new species. X 65. a, front view ; 6, rear 
view; o, apertural view. 
2. Bpiroloculina ewoavata d'Orblgny. X 40. a, front view; b, apertural 

view. 
8. Sigmoilina aaperula (Karrer). X 65. a, front view; 6, apertural view. 
1 Sigmoilina tenitis (Czjzek). X 65. a, front view; b, rear view; o, aper- 
tural view. 

Plate 82. 

FiaL TrUoculina trigonula (Lamarck). X 65. a, front view; b, side view; 
Cf apertural view. 
2. TrUoculina tricarinata d'Orblgny. X 65. a, rear view ; 6, side view ; o, 

apertural view. 
8. TrUoculina bulbosa, new spedes. X 65. a, rear view; b, side view; o, 
apertural view. 

Platb 88. 

F16.I. TrUoouUna projecta, new species. X 65. a, front view; &, rear view; 

c, apertnral view. 
2. BUoculina bulUHdes d'Orblgny. X 65. a, front view; b, side view; 

Cf apertural view, 
a OrbicuHna adunca (Flchtel and Moll). X 80. 



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For explanation of pi ate bee page 86. 



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FOM IXPLANATION OP PLATE BCC PAQC 87. 



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INDEX. 



Page. 

tUir«Tlata, Teztnlarla 61 

atfunet, Orblcollna 84 

Moartoisls, BoUvina i 64 

uqnllateralii, Globlgerlna 67 

agglotliiaiis, TextularU 62 

tmerlcaiia, Tmncatalliia 68 

AmphiftccliH 77 

lesaonll 77 

inomalina, Nonlonlna 74 

uperaU, SlgmoUlDa 81 

anberUna, <)iiliiqiielociillna 79 

bleomli, QDinqaelocQllna 80 

Bicnierliia 56 

nodoaaria 66 

BUocolina 84 

ImUoldea 84 

BolWna <54 

aenaiiensU 64 

cf. B. punctata 64 

robuata-.......— .... .. • 64 

bolbosa, TrOocnllna 88 

bulloldes, BllocuUiia 84 

bttUoldes, Qloblgerina 64 

fflmrlenris, Uylgerlna 62 

earlnata, Teztnlarla 63 

Cbrysalldlna 64 

pulchdla 64 

ClamUna — 67 

commnnls .». 67 

pafifllensls 67 

communis, ClayuUna 67 

eooununls, Nodoearla 69 

concentrlca, Pulvlnullna 71 

conglobata, Globlgeiina 66 

oontorta, <)ulnquelocullna 79 

entlculata, PolystomeUa 77 

crtepa. Polyatomella 76 

CH«tellarla 60 

Itallca 61 

protuberans 61 

rotulata 60 

Taughanl 61 

cQlebrentla, Truncatullna 70 

depretaula, Nonlonlna 72 

Dlicorbela 68 

obtusa 68 

doMa, OloMgerlna 66 

ezcarata, Splrolocullna 84 

fllntll, Gaudrylna 66 

Qaudrylna 66 

fllntU 66 

triangularis 66 

GloMgertna 64 

aequUaAeralls 67 

bulloldes 64 

conglobata 66 



Pi» 

Qloblgerina dubla 66 

Inflata 66 

saccullfera 66 

Inflata, Qloblgerina 66 

Insecta, Nodosarla 69 

Itallca, Crlstellaria 61 

Lagena 68 

striata, Tar. strumosa. 68 

lamlnata, Teztnlarla 62 

leesonll, Amphlsteglna 77 

macella, Polyatomella.. 76 

menardll, Pulvlnullna 71 

Nodoaaria 69 

commfunla . 69 

Inaecta . 69 

raphanlstrum 69 

nodosarla, Blgenerlna 66 

Nonlonlna 72 

anomallna 74 

depressula 72 

panamensls 74 

scapha 78 

obtusa, Dlscorbls 68 

Orblcullna 84 

adunca 84 

Orbullna 67 

unlyersa 67 

panamensls, Nonlonlna 74 

Qulnquelocullna 80 

Teztularla 68 

parislensls, Clayullna 67 

Polyatomella 74 

cratlculata 77 

crispa 76 

macella 76 

sagra 76 

strlato-punctata 74 

projecta, Trllocullna 88 

protuberans, Cristellarla 61 

pulchella, Chrysalldlna 64 

PulTinuUna 70 

concentrica 71 

menardll — 71 

sagra 70 

pygmea, Truncatullna 68 

pygmaea, Uvlgerina 68 

Qulnquelocullna 78 

auberiana 79 

blcomts 80 

contorta 79 

panapensls 80 

■emlnulum 78 

undosa 79 

raphanlstrum, Nodosarla 69 

raphanus. Tar. transrersua, Slphoge- 

nerina 64 

X 



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INDEX. 



reticulata, Slphonlna 72 

robusta, BoUvlna „ 69 

rotnlata, Crlstellarla .... 60 

sacculifera, Globlgerlm — „ 66 

saglttula, Textnlarla ..... 61 

sagra, Polystomella ...... 76 

Polylniilliia. ,„ ■ 70 

scapha, Nonlonlna .... 78 

seminulum, Qnlnqiielocullna. ... 78 

SUpnolllxia 81 

aspemia ^.....i.... 81 

tenuis .... 81 

Siphogenerlna .— .., 64 

raphanua, Tmr. trana- 

versua.....— 64 

Slphonlna .- 72 

reticulata ....... 72 

Splrolocullna ....... 84 

excayata ... 84 

squamosa, YlrguUna ... 68 

striata, var. strumosa, Lagens 68 

strlato-punctata, Polystomella 74 

strumosa, Lagena striata, yar 68 

subagglutlnans, Textularla . 62 

tenuis, Slgmolllna 81 

tenulBtrlata, Uvlgerlna 68 

Textularla 61 

abbrevlata 61 

agglutlnans 62 

carlnata 68 

lamlnata 62 



Faga. 

Textularla panamensls 63 

■aglttula 61 

•Qbagglntlnans 62 

transrersus, Siphogenerlna raphanus, 

var 64 

triangularis, Gaudrylna 66 

trlcarlnata, Trllocullna 82 

trlgonula, Trllocullna 82 

Trllocullna 82 

bulbosa 83 

projecta 88 

trlcarlnata 82 

trlgonula 82 

Truncatullna 68 

amerlcana 68 

culebrensls 70 

pygmea 68 

ungerlana 69 

wuellerstorfl 60 

undosa, Qulnquelocullna 79 

ungerlana, Truncatullna 69 

unlyersa, Orbullna 67 

Uylgerlna 62 

canarlensls 62 

pygmaea 63 

tenulstrlata 63 

yaughanl, Crlstellarla 61 

yirgullna 68 

squamosa 68 

wuellerstorfl, Truncatullna 69 



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SMRBSOHIAM IMSAVIVI'IUN 

UNITED STATES NATIONAL MUSEUM 

Bulletin 103 




CONTRIBtmONS TO THE GEOLOGY AND PALEON* 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



THE LARGER FOSSIL FORAMINIFERA OF 
THE PANAMA CANAL ZONE 



By JOSEPH AUGUSTINE CUSHMAN 
<y tt« Unittd StaUs Gtobgical Sunty 



Extract from Bulletin 103, piget 89-102, «lth Mate* M-49 




WASBINOTON 

OOVBKNMENT PRINTINO OFFICS 

I9U 



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r-r-r ..- x\: i ¥ 



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SMITHSONIAN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 
Bulletin 103 




CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



THE LARGER FOSSIL FORAMINIFERA OF 
THE PANAMA CANAL ZONE 



By JOSEPH AUGUSTINE CUSHMAN 
0} the United Staiei Geological Survey 



Extract from Bulletin 103, pages 89-102, with Platet 34-45 




WASHINGTON 

GOVERNMENT PRINTING OFFrc f 

1918 



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THE LARGER FOSSIL FORAMINIFERA OF THE PANAMA 

CANAL ZONE. 



By Joseph Augustine Cushman, 
Of the United States Geological Survey, 



INTRODUCTION. 

The foraminifera, especially the larger forms of the orbitoids, have 
been little used in America as critical index fossils, except in the 
Yicksburg group; but in Europe, Asia, and the East Lidies they 
have long been used to distinguish horizons. In many geologic 
papers one finds Orhitoides mentioned, probably Orhitoides manteJU 
Morton, and occasionally O, dispcmsus^ O. forhesi, etc. From a crit- 
ical study of the group it soon becomes evident that such identifica- 
tions as have been made of American orbitoids, except those of 
Lemoine and Douvill^, have been largely superficial, and are there- 
fore of little value. Since the earlier work of Giimbel the orbitoid 
foraminifera have with further study been divided largely into the 
four genera Orhitoides^ Orthophvdgminay LepidocycUna^ and MiO' 
gypaina, in general respectively characterizing Cretaceous, Eocene, 
Oligocene, and Miocene formations, but with important exceptions. 
The American forms, with the exception of the work of Lemoine and 
Douvillfi, have not been properly referred to their respective genera, 
although our American Orhitoides manteUi^ described by Morton as 
Nummrdites mantelU in 1833, is the type-species of Lepidocydlincu 
In their work on LepidocycUna Lemoine and Douville ^ describe two 
new American species, L. eaneUei and L. chaperi^ from the Panama 
Canal Zone, figuring also for the first time the critical chambers of 
L. mantelli (Morton). These are all the American species that are 
given, although they call attention to the apparently superficial char- 
acter of the references to Orhitoides in American geologic papers. 
Schlumberger, in his classic works on the genera Orhitoides and 
Orthoph/ragmma^ did not have American material. The American 

1 Sur le Genre LepidocycUna Gfimbel, M6m. Soc. 64ol. France, Pal6ontoIogie, Mem. 82, 
1904. 

89 



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90 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

field is. therefore practically unworked, and the description of our 
species with accurate stratigraphic information is of prime impor- 
tance, as they have been proved elsewhere to be of great use as index 
fossils. 

The collections now in my hands represent the Canal Zone, the 
West Indies, and Coastal Plain Province of the eastern and south- 
em United States. Excellent material was collected in the Panama 
Canal Zone by T. Wayland Vaughan and D. F. MacDonald, and 
is here presented as a beginning in the determination and figuring 
of the American species. This will be followed by papers on the 
West Indian and Coastal Plain species which now, owing to the care- 
ful collecting by Doctor Vaughan and his associates, are represented 
by excellent suites of specimens covering broad ranges, both geo- 
graphically and stratigraphicaUy. As these are gradually worked up 
there will be a mass of data which should be of excellent service in 
the correlation of horizons where these groups are represented, even 
in the absence of Mollusca and other groups of fossils. 

The systematic descriptions of the species of Lepidocyclina^ Nuwr- 
muUtes^ and Orbitolites follow, together with that of a genus and 
species believed to be new, 

LIST OF SPECIES AND THEIR GEOLOGIC OCOURRBNCB. 

Lepidocyclina canellei Lemoine and DouvilW. Oligocene, Culebra 
formation, stations 6019a, Gaillard Cut; 6023, Rio Frijol; 6027, 
Bohio (old station) ; 6891; Bailamons; 6892, 450 feet south of 
switch at Mamei. Also Oligocene of Trinidad. 

Lepidocyclina cTiaperi Lemoine and Douvillfi. Oligocene, Culebra 
formation, stations 6019/, Las Cascadas; 6025, Bohio Bidge 
switch. 

Lepidocyclina vaugJiani^ new species. Oligocene, Emperador lime- 
stone, stations 6021 and 6673, near Caimito Junction ; 6255, half 
mile south of Miraflores Station. 

Lepidocyclina macdonaidi, new species. Oligocene, station 6523, 2 
miles north of David. 

Lepidocyclina panamensis^ new species. Oligocene?, stations 6512, 
river bed, David; Oligocene, 6586^ and 6587, near mouth of 
Tonosi River; probably at 6010, near Miraflores Locks, and 
6012a and 6012^ in Gaillard Cut, in the Culebra formation; 
doubtfully in the Emperador limestone, at station 6015, Empire. 

Lepidocyclina duplicata^ new species. Oligocene, stations 6523, 
2 miles north of David ; and 6586^, near mouth of Tonosi River. 

Heterosfeginoides panamensis^ new si)ecies. Oligocene, Culebra for- 
mation, stations 6011, Gaillard Cut; 6024a, Rio Agua Salud; 
6025, Bohio Ridge switch. Emperador limestone, stations 6015, 
6016, quarries at Empire. 



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6B0L0GY AKD PALEOlTrOLOGY OF THE CANAL ZOISTE. 91 

Orthophragmma rnirmna^ new species, Oligocene ?, station 6612, 

rirer bed, David. 
Nummwlites panamensis^ new species. Oligocene, Culebra forma- 

tioh, stations 6024a, Rio Agua Salud; 6025, Bohio Ridge switch; 

doubtfully at 6026, 2 miles south of Monte Lirio. 
NumanuUtes davidensis^ new species. Oligoc^ie?, stations 6512, river 

bed, David; 6526, Chiriqui. 
Orbitolitea americana^ new species. Oligocene, Culebra formation, 

Gaillard Cut at stations 6013, 60196, and 6020a. 

DESCRIPTIONS OF SPECIES. 

Family NTJMMULITIDAE. 
Genus LEPIDOCYCUNA. 

LEPmOCTCUNA CANELLEI Lemoine iind DootUI^. 

Plate 34, figs. 1-6. 

Lepidocyclina caneUei Lemoine and Douvnxfi, M^m. Soc. G6ol. France, 
Pal6ontologie. Mto. 32, p. 20, pi. 1, fig. 1; pi. 3, fig. 5, 1904, 

Test comparatively small, diameter of largest specimens slightly 
less than four millimeters, thickness a little more than one-fourth the 
diameter; circular in outline, central portion somewhat raised and 
evenly rounded, near the periphery flattened or even slightly con- 
cave; surface in well preserved specimens finely granular or even 
finely papillate, but not strongly so, often appearing smooth to the 
unaided eye. In worn specimens the surface app^ai^ as a series of 
regular hexagonal, honeycomb-like reticulations due to the edges 
of the lateral chambers. 

In vertical section the lateral chambers are seen to be arranged in 
vertical colimms, one directly above the other, from the equatorial 
chambers to the surface, about twelve chambers in each vertical col- 
unm in the central region, the lateral walls hardly thicker than the 
upper or lower surfaces. Chambers of adjacent columns arranged 
alternately, no distinct colmnns present. Equatorial chambers grad* 
ually increasing in size toward the periphery, single throughout, ex- 
tending peripherally beyond the lateral chambers and in surface view 
in well-preserved specimens appearing as a hexagonal reticulation. 
Embryonic chambers nearly equal in size, nearly semicircular in sec- 
tion, their common wall straight. 

Horizontal section showing the equatorial chambers regularly 
hexagonal, those toward the periphery largest. Embryonic cham- 
bers similar to those shown in the vertical view. 

Occurrence. — ^Lemoine and Douvill6 described and figured this 
species from Panama, from Penablanca, also noting it from Mar- 
83701H-18— BuU. 103 7 



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92 BULLETIN 103^ UNITED STATES NATIONAL MUSEUM. 

tinique and Angola. The species from Panama was recorded by 
Dall and by Bagg as Orbitoides forhesi Carpenter.* 

Cat. No. 135216, U.S.N.M., is Lepidocyclina caTieUei Lemoine and 
Douville. Figures 1, 4-6, on plate 34 are from this material, col- 
lected by Hill at Bohio, Panama, where it is very abundant. This 
is the same locality as station 6027 of Vaughan and MacDonald, 
orbitoidal marl, a quarter of a mile northwest of Bohio railroad 
station. In this material L, caneUei is very abundant and makes up 
a considerable proportion of the marl. Parts of five specimens, close 
to one another, are visible in a small part of a section from this 
station. 

Specimens in the collection of the United States National Mu- 
seum, Catalogue No. 107158, from the Oligocene of Trinidad (" Leda 
bed," Naparima) collected by Guppy, are also very evidently Lepido- 
cyclina canellei. 

Specimens of L, caneUei were also very abundant at station 6891, 
foraminiferal limestone from Bailamonas, Canal Zone, collected by 
D. F. MacDonald. 

There is a limestone from station 6892, 450 feet south of switch 
at Mamei, Canal Zone, also collected by MacDonald, which contains 
numerous specimens of a Lepidocyclina in general shape in section 
resembling Z. caneUei^ but the material is very cherty and the finer 
structure is not well preserved. 

A few small weathered specimens from 6019a, Gaillard Cut, oppo- 
site Las Cascadas, seem to belong to this species also; and specimens 
were also obtained at station 6023, along the relocated line of the 
Panama Railroad, at Rio Frijol. The geologic occurrence is in the 
Culebra formation. 

Cat. Nos. 324733-5, U.S.N.M. 

LEPmOCTCUNA CHAPERI Lemoine uid DevrilU. 

Plate 35, figs. 1-3 ; plate 36. 

Lepidocyclina chaperi Lemoine and Douvill6, M6m. Soc. G4ol. France, 
Pal^ontologie, Mto. 32, p. 14. pi. 2, fig. 5, 1904. 

Test of mediiun size, diameter from 8 to 20 millimeters, circular in 
outline, somewhat saddle-shaped, central portion slightly thickened, 
thence gradually and evenly thinning toward the periphery; surface 
where well preserved slightly papillate, usually roughened by ero- 
sion, toward, the periphery often somewhat reticulately depressed 
above the equatorial chambers. 

Vertical section usually curved, lateral chambers numerous, 
breadth much greater than height, columns separated by distinct 
pillars, comparatively few except in the central region where there 

iHill, Geology of Panama, Bull. Mus. Comp. Z5ol., vol. 28, pp. 272, 275, 1897. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 93 

are a few larger than the others ; embryonic chambers of the double 
type, the two chambers nearly equal in size and separated by a 
straight common wall. 

Horizontal section shows similar conditions of the embryonic 
chambers and distinctly hexagonal equatorial chambers. 

Occurrence, — ^Lemoine and Douvillfi described this species from 
Panama (Haut-chagres, San Juan). The figured specimens are from 
United States Geological Survey station 6025, Culebra formation, 
from marl, south end of Bohio Ridge switch, relocated line, Panama 
Railroad, collected by Vaughan and MacDonald. 

Specimens from station 6019-/, Culebra formation, on the west 
side of Gaillard Cut near Las Cascades, seem to represent the micro- 
spheric form of this species. The sections are shown in plate 35, 
figure 3, and plate 36. 

A specimen from station 6526, Chiriqui, Canal Zone, shows a sec- 
tion which from its general proportions strongly suggests Z. cJiaperi, 

Cat. Nos. 324736-8, U.S.N.M. 

LEProOCTCLINA VAUGHANI, new ipeeieik 

Plate 37, lijrs. 1-5 ; plnte 38. 

Test of medium size, 10 millimetei^s or more in diameter, flat, sur- 
face somewhat umbonate in the central portion, gradually sloping to 
the peripheral portion, the outer half of which is nearly flat. Wall 
smooth except for fine papillae. 

Horizontal section shows the peculiarity of the chambers, many of 
which, especially those of the outer peripheral portion are rhomboid, 
those of the inner portion being more typical and hexagonal. These 
are shown especially well' on the sections of the larger specimens, 
those of the smaller specimens showing only the regular hexagonal 
character of the earlier chambers. 

No very good vertical sections were obtained in the thin sections 
but several accidental sections show the characters well. The em- 
bryonic chambers are rather • large, of the usual American type, of 
two nearly equal chambers, lateral chambers in vertical columns 
with a very few, rather well developed pillars. 

Occurrence. — Type-specimen from station 6021, from the Empera- 
dor Limestone in cuttings of the Panama Railroad near Caimito 
Junction, Panama, United States National Museum Catalogue No. 
324739, collected by T. W. Vaughan and D. F. Mac Donald. Speci- 
mens were abundant in this light gray to cream-colored sandy lime- 
stone. Specimens were also abimdant in the collection from the same 
locality collected later by MacDonald under station No. 6673. Speci- 
mens which are apparently the same species are abundant in a fos- 



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94 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

siliferous limy sandstone collected by MacDonald at station No. 
6255 from half a mile south of Miraflores Station on the wagon road 
to Panama. 

LEPmOCTCUNA IfACDONALDI, haw spedM. 

Plate 40, figs. 1-6. 

Test circular, rather small, about 5 to 7 millimeters in diameter, 
thickest in the central region, thence gradually sloping to the 
periphery which for a short distance in from the edge is nearly flat; 
wall rather smooth except the central portion of the umbonal region, 
which has a few pustule-like raised spots at the surface end of the 
vertical pillars. 

Vertical section shows the test widest in the middle, gently sloping 
to near the periphery where the edges are nearly parallel for a short 
distance to the peripheral edge or even slightly increasing in thick- 
ness. Lateral chambers in the central portion in definite vertical 
columns, occasionally slightly overlapping. Equatorial chambers 
not increasing very rapidly in height in megalospheric specimens, 
those at the periphery hardly more than double the height of those 
near the center of the test; embryonic chambers in the megalospheric 
form, large, usually of two nearly equal chambers, but in oblique cut- 
ting these may appear somewhat unequal, plate 40, figures 2 and 3. 

Horizontal sections show chambers somewhat similar to L. 
vauffhani but with the inner half of two walls at nearly right angles, 
the outer wall broadly rounded. The oblique section (pi. 40, fig. 6) 
shows the pillars. 

Occurrence. — ^Type-specimens from station 6523, from orbitoidal 
limestone, 2 miles north of David, Panama^ collected by D. F. Mac- 
Donald, U. S. National Museum Catalogue No. 324740. Specimens 
were abimdant at this station, occurring with L. panamenais and L. 
duplicata. The species were also collected by MacDonald at station 
6512, in the river bed at David. 

LBPmOCTCLINA PANAMENSI8, new spMiM. 

Plate 39. flgs. 1-6 ; plate 42. 

Test circular, small, central portion very strongly umbonate, thick, 
rapidly decreasing in thickness peripherally, the peripheral portion 
thin and flattened, the raised central portion only one-third to one- 
fifth the entire diameter, which ranges from three to six millimeters; 
occasional specimens, perhaps representing the microspheric form, 
up to 10 or 12 millimeters in diameter; surface smooth except for 
the unbonal portion which has a few large pustule-like projections 
marking the ends of the internal pillars. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 95 

The vertical section shows very peculiar embryonic chambers in 
that they do not exhibit the usual charactei^ of American species, but 
have a broad and much flattened central chamber two to four times 
as broad as high with a compressed, partially encircling chamber, 
which in section is usually cut on the opposite sides of the central 
chamber. In some cases there seems to be an irregular mass of three 
or four more or less nearly spherical chambers. In the former case 
these central chambers in section are nearly as wide as the whole 
umbonal portion of the test. Lateral chambers, usually about twice 
as wide as high, the outer wall often somewhat arched toward the 
exterior of the test, arranged in vertical columns. Pillars not dis- 
tinct except in the central portion where there are a few strong ones 
increasing rather rapidly in diameter toward the periphery, usually 
about 9 or 10 chambers in a vertical column in the center of the 
umbilical region. The peripheral region has only a thin coating of 
lateral chambers, the last formed layer present only on the outer 
half of the periphery and often none at all present on the last quar- 
ter of the test toward the periphery, the surface being made up by 
upper and lower walls of the equatorial chambers. Equatorial cham- 
bers nimierous, comparatively broad, the peripheral wall convex out- 
wardly toward the periphery, the chambers at least as wide as high. 

In horizontal section the equatorial chambers are usually some- 
what irregularly hexagonal near the center, toward the periphery 
more or less rhomboid with the outer peripheral wall curved. 

As far as described material is concerned this is an unusual form 
for American species of LepidocycUna, especially in its embryonic 
chambers. 

Occurrence. — Type-specimen, vertical sections, U. S. National 
Museum Catalogue No. 324741. The species is fairly abundant at 
stations GoSQe and 6587 from near the mouth of Tonosi River, 
Panama, D. F. MacDonald, collector. It was also collected by Mac- 
Donald at station 6512, river bed, David.. 

At stations 6010, 600 or 700 feet south of the Miraflores Locks, and 
6012a and 6012r, south of Empire Bridge, in the Culebra formation, 
specimens of small orbitoids occur, but they are not sufficiently well 
presen-ed for positive identification. Although those from the latter 
station seem somewhat like Z. pananianensis in their thin borders 
and raised center w^ith papillae, they can not be specifically identified 
with certainty. At other stations poorly preserved orbitoid fo- 
raminifera occur, but their specific identity can not be accurately 
determined. Specimens doubtfully referable to Z. panamensis were 
obtained in the Emperador limestone, at station 6015, Empire. 



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96 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

MULTICYCLINA, new subgenus. 

Subgenus differing from typical Lepidocyclina in the equatorial 
chambers which instead of being in a single series become complex 
toward the periphery and may consist of several series. 

Type of. the suhgenus. — Lepidocyclina duplicata Cushman. 

LEPIDOCTLINA DUPLICATA, new ipedef. 

Plate 41, figs. 2-4. 

Test of medium size, 10 to 14 millimeters in diameter, very much 
thickened in the umbonal region, usually the thickness about one-half 
the diameter; without the flattened periphery the central portion 
is subspherical, thinning rapidly toward the periphery, then thick- 
ening again at the margin, which is often doubly plicate in the best 
preserved specimens. Surface of the umbonal portion studded with 
numerous fine papillae marking the surface terminations of the pil- 
lars, peripheral portion nearly smooth. 

^>rtical section showing the embryonic chambers as very small, 
apparently microspheric in the specimens sectioned, appearing spiral 
as is usual in the microspheric form. Lateral chambers numerous, 
flattened or lenticular, the numerous pillars as wide as or wider than 
the intermediate columns of chambers, especially in the central por- 
tion, rapidly increasing in size toward the surface. Equatorial 
chambers very small near the center, gi'adually increasing in size 
toward the periphery where they become multiple instead of single 
as is usually the case, and make three or four vertical scries, each 
with numerous fine apertural pores on the outer convex wall. 

Horizontal section shows the increase in size of the equatorial 
chambers which toward the center seem hexagonal and toward the 
periphery almost rhomboid with the outer half convex. 

Of somewhat similar character as far as the duplication of equa- 
toral chambers is the species described by Martin from Java, Z, 
multipartita (Martin), and the form described by Jones and Parker 
from Christmas Island, L. i7v<ulae nafalis^ var. inequalis (Jones and 
Parker). 

Occurrence, — Type-specimen, U. S. National Museum, Catalogue 
No. 324742. Specimens were very abundant, weathered out of an or- 
bitoid limestone, at Station 6523, 2 miles north of David, Panama, 
D. F. MacDonald, collector. They also occur with other species of 
Lepidocyclina at Station 6586^ from near the mouth of Tonosi River, 
D. F. MacDonald, collector. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 97 

HETEROSTEGINOIDES, new geniuu 

Test generally lenticular, somewhat excentric, one side extended 
peripherally more than the other, chambers rather coarsely per- 
forate, embryonic chambers, often two, of nearly equal size, thick 
w ailed, chambers added as in Heterostegina, in a revolving series ex- 
tending from the umbonal region on both sides to the periphery, 
chambers hemispherical, the outer side strongly convex and all 
coarsely perforate, the equatorial chambers larger than the lateral 
ones and nearly spherical. 

Type of the genus. — Ileterosteginoides panamensisy new species. 

HETEROSTEGINOroES PANAMENSIS. mw spedM. 

Plate 43, figs. 1-8. 

Test biconvex, somewhat more strongly convex on one side than on 
the other, revolving edge indistinct, surface unevenly rugose, or ir- 
regularly pustulate, thickest in the umbonal region. Vertical section 
showing the embryonic chambers as an equal pair of nearly spherical, 
thick-walled chambers, equatorial chambers also nearly spherical, 
lateral chambers hemispherical with the curved side outermost, ir- 
regularly piled above the equatorial chambers. Horizontal section 
showing the central chambers with the equatorial chambers arranged 
in an irregular semi-spiral manner about them. 

Test small, between 1 and 2 millimeters in diameter. Cat. Nos. 
324743-4, U.S.N.M. 

Occurrence. — This species was abundant in the Culebra formation 
at station 6025, from marl, south end of Bohio Ridge switch, relo- 
cated line, Panama Railroad, collected by Vaughan and MacDonald. 
There are also numerous specimens at station 6011, Culebra forma- 
tion, along east side of Gaillard Cut, collected by Vaughan and Mac- 
Donald. It was also collected in the Culebra formation at station 
6024r-«, Rio Agua Salud, and is doubtfully present in the Emperador 
limestone at stations 6015 and 6016, in Empire. 

This species, which in external appearance somewhat resembles a 
small orbitoid or nummulite, may be distinguished from most species 
of either group by its comparatively coarse pustulate exterior. In 
section it can easily be recognized by its peculiar structure. 

Genus ORTHOPHRAGMINA. 

ORTHOPHBAGIONA MINniA, imw tpedes. 
Plate 41, fig. 1. 

Test circular, very small, slightly more than 2 millimeters in 
diameter^ thickness somewhat less than half the diameter, central 
portion very strongly umbonate, evenly rounded to a point about 



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98 BULLETIN 103, UNITED STATES NAJDIONAL MUSEUM. 

two-thirds of the distance from the center to the periphery, from 
which point to the periphery the surface is nearly fiat; surface of 
the test comparatively smooth. 

Horizontal section through the equatorial chambers shows very 
fine rectangular chambers and the embryonic chambers nearly equal 
in size. 

Vertical section (fig. 1) shows well the contour of the test in this 
section, the strongly curved central umbonate portion making up 
two-thirds or more of the width and the peripheral flange with its 
nearly parallel sides. The chambers are very small, except the 
embryonic central chambers, which are nearly equal and have a 
straight division line between them. The lateral chambers are in 
vertical columns, but the test is without pillars. In the central region 
there may be more than 20 chambers in a vertical column, and even 
on the peripheral flange there are usually three or four in a column 
on each side of the equatorial chambers. 

Occurrence. — ^Type-specimen — the vertical section here figured. 
Collection of the U. S. National Museum Catalogue No. 324745. 

The species is abundant at United States Geological Survey sta- 
tion 6512 in the white limestone, in the river bed above the ice plant 
near David, Panama, collected by D. F. MacDonald. 

This is a very small species yet it has an abundance of very fine 
chambers. There is an exceptional development of lateral chambers 
in the region of the periphery. 

Genus NUMMULITES. 

NUMBfUUTES PANAMENSIS. new specie*. 

Plate 43, figs. 9, 10. 

Test small, about 1^ millimeters in diameter, much compressed, 
chambers very numerous, about 22 in the last formed coil, each in 
section two to three times as high as long, test of about four whorls, 
walls comparatively thick, whole test lenticular, peripheral margin 
broadly rounded, central portion nearly flat. 

Occurrence. — Specimens occur with some frequency in the Culebra 
formation at station 6025, in marl, south end of Bohio Ridge switch, 
relocated line, Panama Railroad, collected by Vaughan and Mac- 
Donald. Type-specimen, U. S. National Museum Catalogue No. 
324746. The species was also collected in the Culebra formation at 
station 6024-a, Rio Agua Salud, and doubtfully at station 6026, 2 
miles south of Monte Lirio, on the relocated line of the Panama 
Railroad. 

NUMMULITES DAVIDBNSIS. new spedei. 

Plate 43, fig. U. 

Test comparatively small, about 3^ millimeters in diameter, some- 
what compressed, chambers about twice as high as long in median 



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QBOLOGY Ain> PALEONTOLOGY OF THE OANAL ZONB. 99 

sections, test of three or four whorls, walls of medium thickness, 
the upright wall rather strongly recurved backward in the central 
portion, 15 or 16 chambers in the last formed whorl. 

Occtkrrenoe. — Specimens were not common at station 6512 from 
white limestone in river bed above ice plant near David, Panama, 
collected by D. F. MacDonald. 

Type-specimen.— CBi. No. 324751, U.S.N.M. 

In material from station 6526 from Chiriqui, Canal Zone, col- 
lected by MacDonald, numerous specimens occur which in section 
seem identical with this species. 



Family MIUOLIDAE. 

OBSnOLITES AMERICANA, new ipmIm. 

Plate 43, figs. 12-14 ; plate 44, figs. 1, 2 ; plate 4^. 



T j^J^/oBBrtoiJTJ 



Test flat, of medium size, larger specimens about 10 millimeters 
in diameter, chambers with the outer wall strongly convex, the inner 
wall running backward and bluntly pointed, side walls parallel, 
chambers in two or more tiers ; tests mostly microspheric, one (pi. 43, 
fig. 14) apparently megalospheric, and one (pi. 44, fig. 2) seemingly 
originating from a fragment of an older test. Numerous specimens, 
especially plate 45, figure 1, show evidence of breakage and repair. 
The apertures between the chambers are not evident, as the material 
largely consists of internal casts of the chambers. Some of the 
specimens suggest the genus Presorites of the Cretaceous described 
by DouvilM, but this is probably due to the condition of fossilization. 

Occurrence, — Specimens which seem referable to this species are 
from the following stations at Panama, collected by Vaughan and 
MacDonald: Culebra formation, 6013, east side of Ghdllard Cut; 
60196, 6019-€--/, west side of GaiUard Cut near Las Cascadas; and 
6020a-<? of the same section. Also collected in the Emperadoi; lime- 
stone at station 6015, in Empire. 

Type-specimen.— Csit. No. 324748, U.S.N.M., from station 6020a. 

EXPLANATION OP PLATES. 
PlATE 84. 

Lepidocyclina oanellei Lemolne and DoayUl& 

Fkft. 1. View of exterior of spedmen X 10, a portion of a second specimen show- 
ing above the first, from Bohio, Panama. (n.S.NJ!ii. Oat No. 185216.) 



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100 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

2. Horizontal section X 10, showing embryonic cliambers and liexagonal 

equatorial chambers, from west side of Gaillard Cut near Las Cas- 
cadas (U.S.G.S. station 6019a). 

3. Horizontal section showing hexagonal equatorial chambers and irregu- 

larities in the annuli due to repairs of breakage, a X 10; 6 X 20. 
Same locality as No. 2 above. 

4. Slightly oblique section X 20, showing narrow zone of equatorial cham- 

bers and two broader zones of lateral chambers, the latter with a very 
evident lack of pillars. Same locality as No. 1 above. 

5. Vertical section at one side of embryonic chambers showing general 

characters of equatorial and lateral chambers X 20. Same locality 
as No. 1 above. 

6. Vertical section through the embryonic chambers showing the two 

nearly equal chambers with the straight wall dividing tlie two, X 20. 
Same locality as No. 1 above. 

PLATk: 35. 

Lepidocyclina chaperi Lemoine and Douviil^. 

Fig. 1. Exterior view of specimen X 5. Specimen broken. From upper part 
of Culebra formation, from Panama Railroad, southern switch, Bohlo 
Ridge, in light-colored limy sandstone (U.S.G.S. station 6025). 

2. Exterior view of small, more complete specimen from same locality as 

the preceding, X 5. 

3. Horizontal section showing early chambers of the microspheric form of 

the species. X 20. From west ^Ide of Gaillard Cut near Las Cascadas 
(U.S.G.S. station 6019/). . 

Plate 36. 

Lepidocyclina chaperi Lemoine and Douvill^. 

Horizontal section X 10, showing early central chambers and hexagonal 
chambers of the equatorial region (U.S.G.S. station 6019f). 

Plate 37. 

Lepidocyclina vaughani^ new species. 

Fig. 1. View of exterior of specimen X 5, with flat periphery and umbonate 
center, from limy sandstone half a mile south of Miraflores Station, 
on wagon road to Panama (station 6255). 

2. Horizontal section of young specimen with regularly hexagonal equa- 

torial chambers X 20 (same locality as No. 1). 

3. Oblique section X 20, with narrow zone of regularly hexagonal equa- 

torial chambers and broader zones of lateral chambers and a straight 
division wall (same locality as No. 1). 

4. Specimen showing zone of equatorial chambers about peripheral por- 

tion, lateral chambers covering them In the center X 10. From lime- 
stone in cut of relocated line of Panama Railroad opposite San Pablo 
and about 4 miles north of Gamboa bridge (station 6673). 

5. Portion of vertical section (slightly oblique) through the embryonic 

chambers, X 20 (same locality as No. 1). 



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OBOLOGY AKD PALBONTOLOaY OF THB OAKAL ZONB. 101 

Plate 38. 

Lepidooyclina vaughani, new species. 

Specimen X 20, showing peripheral zone cut through the equatorial chambers 
and central portion covered by lateral chambers. From limestone in cut of re- 
located line of Panama Railroad opposite San Pablo and about 4 miles north 
of Gamboa bridge (station 6673). 

Plate 39. 

LcpidoQyclitia panamensis, new species. 

Fig. 1. Very young specimen In vertical section consisting of embryonic cham- 
bers and one or two following chambers, X 20. 
2-4. Vertical sections with broad embryonic chambers and showing the rela- 
tion of equatorial and lateral chambers, vertical columns of lateral 
chambers with intermediate pillars, X 20. 

5. Oblique section with zone of hexagonal equatorial chambers, X 20. 

6. Section of rock with six specimens lying closely adjacent, four of these 

cut through the embryonic chambers, X 20. 
All eqpecimens from near the mouth of Tonosi River, Panama (station 
6586e). 

Plate 40. 

Lepidooyclina macdonaldi, new species. 

Fig- 1. Exterior view of specimen, X 10, showing pillars appearing at the 
surface as raised area. 
2-5. Vertical sections (slightly oblique) through the embryonic chambers, 
which when cut in plane at right angles to division wall show nearly 
equal chambers with the division wall straight or very slightly 
curved. Pillars evident, especially in Nos. 2 and 5. X 20. 
6. Oblique section, X 20, showing zone of "lozenge-shaped" equatorial 
chambers with lateral chambers on each side. The upper series 
showing the cut sections of pillars. 
All specimens from limestone 2 miles north of David, Panama (station 6623). 

Plate 41. 

Orihophragmina minima, new species. 

Fig. 1. Vertical section, X 20, showing general outline and numerous very fine 
chambers. From white limestone in river bed above ice plant, David, 
Panama (U.S.G.S. station 6512). 

Lepidocyclina duplicata, new species. 

2. Exterior view of type, X 5, showing raised center and depressed area 

inside the raised periphery. 

3. Portions of vertical section showing great increase In width of equa- 

torial zone, multiplication of chambers toward the periphery, heavy 
pillars and wide lateral chambers. X 20. 



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102 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

4. Portion of oblique section showing narrow zone of " lozenge-shi^ted ** 
equatorial chambers, perforations of peripheral wall of outer equa- 
torial cliambers and perforated pillars among the lateral diambers. 
X 20. 
All specimens of L. dupHoata from limestone, 2 miles north of David, Panama 
(station 6528). 

PLATB42. 

Section of limestone from station 6523, 2 miles north of David, flowing nu- 
merous specimens of Lepidocyclina, X 20. Left center, L. panamenHs ¥rith 
broad embryonic chambers ; lower middle L. macdonaldi with subspherical em- 
bryonic chambers ; at right a portion of L. duplicaia, 

Plate 43. 
Heterosteginoides panamenaiSf new genus and new species. 

Fig. 1, 2. External view of [Q>ecimens, X 10, from limy sandstone, east side of 

Gaillard Cut (station 6011). 
3-6. Vertical portions, X 20, showing irregular piling of lateral chambers ; 

fig. 6 with two embryonic chambers with thick walls. Specimens 

from limy sandstone near southern switch, Bohio Ridge (station 

6026). 
7, 8. Horizontal sections, X 20, from same locality at Bohio. 

NummuHtea panamensiSy new i^)ecies. 

9. Horizontal section, X 20, from limy sandstone near southern switch* 

Bohio Ridge (station 6025). 
10. Vertical section from same roclc specimen, X 10. 

NummuUtes davidensis, new species. 

IL Horizontal section, X 20, from white limestone in river bed above ice 
plant, David, Panama (station 6512). 

OrhitoUtes amerioana, new Q)ecieB. 

12-14. Horizontal sections, X 10, specimens from west side of Gaillard Gut, 
near Las Cascadas (station 6020a). 

PLATB44. 

OrbitoHte$ amerioana, new species. 

Fig. 1. Horizontal section of large specimen, X 10, from Gaillard Cut, near Las 
Cascadas (station 6019-5). 
2. Horizontal section, brolcen, showing two layers of chambers and inslda 
cast of outer surface, X 20 (station 6020a). 

Platb45. 

Orbitolites americana, new species. 

Specimen in horizontal section, X 20, showing several areas of breakage and 
subsequent repair, shown by the angular reentrants of the annul! in varloiis 
places. From Gaillard Cut, near Las Cascadas (station 6020a). 



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BULLETIN 103 PL. 34 




Larger Fossil Foraminifera from Panama. 

For eXPLANATION or PLArB SBK PAOes 89-100. 



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BULLETIN 103 PL. 86 




Larger Fossil Foraminifera from Panama. 

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BULLETIN 103 PL. 36 




Larger Fossil Foraminifera from Panama. 

For explanation of plate tEi page 1 00. 



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BULLETIN 103 PL. 37 




LARGER Fossil Foraminifera from Panama. 

For explamatiom of plate see paoe 100' 



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BULLETIN 103 PL. 38 




Larger Fossil Foraminifera from Panama. 

For explanation op platc see page IOi. 



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BULLETIN 103 PL. 89 




Larger Fossil Foraminifera from Panama. 

For exPLANATioN op plate tee paqc 10 1 



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BULLETIN 103 PL. 40 




Larger Fossil Foraminifera from Panama. 

For kxplanation op platk skc pack ioi. 



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BULLETIN 103 PL. 40 




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For eXPLANATION op PLATE tEB PAQB 101. 



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BULLETIN 103 PL. 41 




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For cxplanation op plate vce pages 1 01. los. 



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LARGER Fossil Foraminifera from Panama. 

For eXPLANATION of PLATK tCE PAQK 102. 



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BULLETIN 103 PL. 43 




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BULLETIN 103 PL. 44 




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BULLETIN 103 PL. 46 




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INDEX. 



•mericana* Orbitolitet .. 99 

ctnellel, IjepMocydlna,,, .,„».->« 91 

cbaperl* Ijepldocycllna .— . 92 

dATidenslB, NammiiliteB ....^ 98 

dapUcata* liepidocydlna 96 

Heteroateglnoldea . 97 

panamenslfl 97 

LeiildocycUiia ... 91 

canellei.. 91 

cbaperl 92 

dnpUcata . .- 90 

macdonaldl ...— — 94 

panamenels ....... 94 

▼aoghani .... 98 



macdonaldl, Lepldocyellna 94 

minima, Orthophragmlna .. .. 97 

Mnltlcydlna 96 

NnmmnllteB 98 

davldensls 98 

panamensto 98 

Orbltolltet 99 

amerlcana. . 99 

Orthophragmlna .. 97 

minima 99 

panamensls, Heteroateglnoldes 97 

panamenslB, Lepldocyellna 94 

panam^isls, Nnmmnlltet 98 

^nghanl, Lepldocyellna 98 

K 



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SMITHSONIAN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 
Bulletin 103 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



FOSSIL ECHINI OF THE PANAMA CANAL 
ZONE AND COSTA RICA 



By ROBERT TRACY JACKSON 
0} Peterborough, New Hampshire 



Extract from Bulletin 103, pages 103-116, with Plates 46-52 




WASHINGTON 

GOVERNMENT PRINTING OFHCE 

1918 



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-FOSSIL F.CHINI OF THE PANAMA CANAL ZONE AND 

COSTA RICA. 



By Robert Tract Jackson, 
Of Peterborough, New Hampshire. 



INTRODUCTION. 

The following is essentially a reprint of my paper bearing the 
same title published in the Proceedings of the United States National 
Museum, volume 53, pages 489-501, plates 62-68, September 24, 1917: 

The fossil echini of the Panama Canal Zone were submitted to me 
for study and description by Dr. T. Wayland Vaughan as part of the 
studies he is making in that region in connection with his investiga- 
tions of the geology of the Costal Plain of the United States and of 
the West Indies. The material oJtitains some very interesting 
species, particularly in the genus Encope^ of which there are three 
•new forms. Some of the material is well preserved, and parts are 
fragmentary. A number of specimens too poorly preserved, or too 
fragmentary for specific determination, indicate that a more ex- 
tensive echinoid f aima may be found by further search. 

I wish to express my heartiest thanks to my friend. Dr. Hubert 
Lyman Clark, of the Museum of Comparative Zoology, who, with his 
great knowledge of Clypeastroids and Spatangoids, helped me 
materially in preparing this report. 

LIST OF SPECIES AND THEIR GEOLOGIC OCCURRENCE. 

Clypeaster lanceolatus Cotteau. Upper Oligocene, Emperador lime- 
stone, Gaillard Cut, stations 58666, 6671. 

Clypeaster gatuni Jackson. Miocene,^ Gatun formation, station 5662, 
near Gatun Dam site ; and at station 6237, north of Ancon Hill, 
about 4 miles south of Diablo ridge. 

Encope annectans Jackson.^ Miocene, Gatun formation, station 
5846, Spillway, Gatun Dam. 

Encope platytata Jackson. Miocene,* Gatun formation, station 
6029a, one-quarter to one-half mile from Camp Cotton, toward 
Monte Lirio. 

^ This formation is more appropriately referable to the lower Miocene, 1. e., Bordigalianr 
tban to the npper Oligocene. — T. W. V. 

108 



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104 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Encope megatrema Jackson. Miocene,^ Gatun formation, station 
6030, about one and one-half miles from Camp Cotton, toward 
Monte Lirio. 

Echinolcmipas aerrdorhis Guppy. Upper Oligocene, Emperador lime- 
stone, Gaillard Cut, stations 58666 and 6019g^. 

Schisaster amdge)' W. B. Clark. Miocene ( ?),^ Bonilla, Costa Rica. 

SchiBoster eristatus Jackson. Miocene (?),* Brazil, Costa Bica, sta- 
tion 5505. 

Schizaster panamensis Jackson. Miocene,* Gatun formation, near 
Gatun, at stations 6008 and 7294. 

DESCRIPTION OF SPECIES. 

CLTPEASTER LANCEOLATUS Cottean. 

Plate 46, flga 1, 2. 

Clypeaater lanceolatua, Cotteau, Descripclon de los Equlnoldes Fossiles de la 
Isla de Cuba, Bol. Com. del. Mapa Geologlco de Espana, vol. 22, 1897, p. 
39, pL 9, figs. 1, 2, 3.-— Jackson, Ptoc. U. S. Nat Mus., vol. 53, 1917, p. 
490, pi. 62, figs. 1, 2. 

This species is one of the fe^ in the series from the Panama Canal 
Zone that seems referable to an already published species. There 
are seven specimens, all in good condition of preservation and repre- 
senting two localities which, however, from the character of the 
material may be nearly associated. I give measurements of the 
largest specimen of the set. Length, 95 mm. ; width, 77inm. ; height, 
21 mm. Test elongate, wider behind than in front, moderately ele- 
vated, deeply concave in ventral view. Ambulacral petals elevated, 
distally acuminate, nearly closed and pinched up as if squeezed 
between the thumb and finger. Anterior petal III equal in length 
to petals I and V and a few millimeters longer than are the anterior 
pair II and IV. The anterior petal III is more widely separated from 
petals II and IV than are those latter from I and V. Interporiferous 
areas of petals are elevated, wide, being about equal to both porif- 
erous areas. Interambulacra are narrow, extremely so near the 
apical disk. Tubercles are small and of about the same size dorsally 
and ventrally. Apical disk is central, mouth central, deeply sunken, 
periproct ventral, about four mm. from the posterior border of the 
test. The original material described by Cotteau is from the " Mio- 
cene " of Matanzas, Cuba, where he says it is very rare. It is appar- 
ently more or less common in the Canal Zone, as there are seven speci- 
mens from that region. 

1 This formatioii Is more appropriately referable to the lower Miocene, 1. e., Bnrdlffallan, 
than to the Upper Oligocene. — T. W. V. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 105 

Localities and geologic occwrrence. — ^Upper Oligocene; Emperador 
limestone. Upper Limestone, Las Cascadas, Panama, D. F. Mac- 
Donald, collector, U. S. National Museum station No. 6671, two 
specimens, U. S. Nat. Mus. Cat. No. 324452 ; also Panama Canal Zone, 
upper Limestone bed, near Tower " N " (opposite Las Cascadas, Gail- 
lard Cut) D. F. MacDonald, collector, 1911, U. S. National Museum 
station No. 5866 6, five specimens, U. S. Nat. Mus. Cat. No. 324451. 

CLTPEASTER GATUNI Jackson. 

Plate 47, fig. 1 ; plate 48, fig. 1. 
Clypeaster gatuni Jackson, Proc. U. S. Nat. Mus., vol. 53, 1917, p. 491, pi. 
63, fig. 1; pi. 64, fig. 1. 

This species is represented by a fine, large specimen in perfect con- 
dition of preservation. Two additional specimens much worn and 
incomplete are also referred to it. 

The type measures 146 mm. in length, 122 nmi. in width, and 35 
mm. in height. The test is elongate, moderately pentagonal in out- 
line, with slight incurving of the borders in interambulacral areas 
1, 2, 3, and 4. Its greatest width is across ambulacra II and IV. 
Ventrally the test is deeply concave, being flat only on the border. 
The ambulacral petal III is equal in length to petals I and V and 
a few millimeters longer than are petals II and IV. The petals are 
equidistant, highly elevated, and open at their distal ends. Ventrally, 
five deep ambulacral grooves extend to the mouth. Interambu- 
lacra are broad on the border of the test, narrowing up dorsally and 
very narrow near the apical disk. Each of the interambulacra 
between the petals are strongly elevated as if pinched up. The 
apical disk is slightly anterior to the middle of the test and is very 
small The mouth is central, deeply sunken. The periproct is 
ventral, slightly elliptical, its posterior border 5 mm. from the 
posterior limits of the test. Tubercles are small, covering the 
dorsal surface of the test, ventrally the same, but slightly larger. 

Clypeaster gatuni approaches nearest, perhaps, to C. bowersi Weaver, 
but diflFers in the shape of the test, the deeply concave base, the 
shape and proportionate size of the petals and interambulacra dor- 
sally, and the fact that the periproct is ventral instead of terminal. 

Locality and geologic occurrence, — Gatun formation, Miocene. 
Panama Canal Zone, near Gatun Dam site, D. F. MacDonald, col- 
lector, 1911, holotype, U. S. National Museum, station No. 5662, 
one specimen. 

Limestone in swamp, north of Ancon Hill, about 4 miles south of 
Diablo Ridge in the upper Oligocene Emperador limestone, IT. S. 
National Museum, station No. 6237, two specimens. 

Holotype.— C^t. No. 824458, U.S.NJt 

This species is present on both the Atlantic and Pacific sides of 
the Isthmus. 



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106 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

ENCOPB ANNECTANS JackMn. 

Plate 49, figs. 1, 2 ; plate 50. fig. 1. 

Encope annecians Jackson, Proc. U. S. Nat Mus., vol. 53, 1917, p. 491, text 
fig. 1; pi. 65, flga 1, 2; pi. 66, fig. 1. 

This interesting species is represented by three specimens which 
include two tests free from matrix and more or less complete, and 
a sandstone mould of the exterior of the ventral side of a specimen 
which is the largest of the three. 




Fig. 1. — Encope annectans. Drawing op the typb-specimbn, natural size, 
parts ark indicated by dotted lines. 



Kkstobbd 



In shape, the specimens are thin, flattened, and nearly circular in 
outline, excepting for the reentrant marginal ambulacral notches. 
The edges are thin, exceptionally so for the genus, and the whole 
test superficially is scutelliform. In the anterior ambulacrum III 
there is a shallow rounded notch, and in the lateral ambulacra are 
deeper and narrower notches, the deepest being in the posterior pair 
of ambulacra, IV and V. The apical disk is central. The peristome 
is small and also central. Continuing posteriorly from the peri- 
stome on the ventral side is a quite deep groove, and on the dorsal 
side is a shorter and shallower groove. These grooves do not form 
a hole through the test, but represent the incipient beginnings of the 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 107 

lunule which is characteristic in Encope of the posterior interambula- 
cruin 5. This is most interesting and is discussed later at length. 

The type-specimen measures 86 mnL in length, 89 mm. in width, 
and 8 mm. in height. The highest point is distinctly anterior to the 
middle of the test. The specimen represented by a mould of the 
ventral side is somewhat larger than the type. It measures about 
93 mm. in length by about 96 nun. in width. 

In the type, the ambulacral petals are broad, about equal in length 
in the several areas, the posterior pair extending back to a line with 
the anterior limit of the lunule in interambulacrum 6. Ventrally, the 
ambulacral furrows are deep, slightly curved outward from the 
median line of each ambulacrum, forking near the border of the test, 
each furrow giving off a forked branch at nearly a right angle to the 
main furrow. The apical disk is central, but details are obscured 
owing to local imperfections in both specimens showing the dorsal 
side. Interambulacral areas are narrow in the petaloid areas, wide 
near the margin of the test The whole dorsal surface of the test is 
covered with small tubercles; on the ventral side of the test the 
tubercles are somewhat larger, but they are reduced in size or want- 
ing along the lines of the ambulacral furrows. The mouth is small 
and central in position. The periproct is small, oval in outline, and 
situated at nearly one-third the distance from the mouth to posterior 
border of the test. 

The lunule of interambulacrum 6 is the remarkable and most in- 
teresting feature of this species. Ventrally, it consists of an im- 
pressed area 15 mm. long by 2 mm. wide, extending to and being 
confluent with the opening of the periproct. Dorsally, the lunule 
also consists of an impressed area lying above the middle of the ven- 
tral lunular depression and measuring 10 mm. in length by 2 mm. in 
width. This is the only species in the genus recorded in which the 
lunule fails to make an opening through the test. Structually, it 
is most interesting, as it closely resembles the condition in a young 
specimen of Mellita aexiesperforata (Leske) from the west coast of 
Florida, 30 fathoms, No. 2900, Museum of Comparative Zoology. 
This yoimg MeUita^ which measures 9 mm. in length, has no notches 
or lunules as yet developed in the ambulacral areas, but in interam- 
bulacrum 5, as viewed ventrally, there is a distinct impressed area 
marking the initial beginnings of a lunule as in our specimen of adult 
Encope annecfans. It should be stated that this specimen of Mellita 
is probably exceptional in holding this youthful character so late, as 
in a small series of younger specimens of M. sexiesperforata measur- 
ing from 4 to 7 mm. in length, all have a perforate lunule in interam- 
bulacrum 5. This latter set is from Salt Key, Bahamas, No. 2439, 
Museum of Comparative Zoology. As pointed out by Mr. Agassiz 
(Revision of the Echini, pp. 320-324) in Mellita aexiesperforata^ the 
8370^^—18— BuU. 103 8 



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108 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

ambulacral and int^rambnlacral lunules develop by resorption through 
the test, whereas in the other species of MeUita^ as far as known, the 
ambulacral lunules are developed by the inclusion of marginal 
notches and the interambidacral lunule alone is formed by invagina- 
tion through the test. 

Encope armectans is primitive like the other fossil species of Encope 
in that the ambidacral notches are not inclosed to form lunules but 
are still shallow and open. It is undoubtedly the most primitiye of 
the genus in that the lunide in interambulacrum 5 is still imper- 
forate. It makes an approach to the Becent Encope fwichelim 
Agassiz of the Gidf of Mexico and E. grandia Agassiz of the Gulf of 
California which are the only living species characterized by open 
marginal notches. On the other hand, E. cmnectans resembles En- 
cope micropora Agassiz of the West Coast in the form of the test 
and the position of the interambulacral lunule. 

Locality and geologic occurrence. — Gatun formation, Miocene, 
Panama Canal Zone, Spillway at Gatun Dam site, D. F. MacDonald, 
collector, U. S. National Museum station No. 6846, three specimens. 

Type.— C9it. No. 324454, U.S.N.M. Paratype.—Czt. No. 324466, 
U.S.N.M. 

ENCOPE PLATYTATA Jackflon. 

Plate 51. figs. 1, 2. 

Encope platytata Jackson, Proc. U. S. Nat. Mus., vol. 53, 1917, p. 494, text fig. 
2; pi. 67, ngs. 1,2. 

There is only a single specimen representing this species, and 
while it is imperfect, it yet has the essential parts preserved that are 
necessary for a description. As in the last described species, E. an- 
nectans^ this species, E. platytata^ is thin, flattened, and if complete, 
apparently would be nearly circular in outline excepting for the am- 
bulacral notches. If complete, the specimen woidd measure as esti- 
mated about 100 mm. in length and 100 mm. in width. The greatest 
height of the test is in the apical region, where it measures 10 mm. 
As the ventral side of the test is somewhat concave instead of being 
flat, the thickness of the test at the center, as measured by calipers, 
is somewhat less than the height and measures only 8 mm. 

The anterior ambulacral notch of area III is very shallow and 
rounded. The notches of the lateral anterior ambulacra II and IV 
are also rounded but deeper than the notch of area III. Presumably 
the notches of the posterior ambulacra I and V, if preserved, would 
be similar but somewhat deeper, as this is the usual character in 
associated species. The lunule in interambulacrum 5 is small, but 
passes directly through the test instead of being imperforate as in 
Encope annectans. This lunule is only preserved for the anterior 
part of its extent as shown in the figures. The mouth is small and 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 



109 



central in position, the periproct is elongate oval, its anterior border 
is 13 mm. posterior to the border of the mouth opening. Poste- 
riorly the periproct is confluent with the infolded depression of the 
interambulacral limule. 

The ambulacral petals are rather narrow in this specimen, measur- 
ing 13 mm. in width. The odd anterior ambulacral petal is longer 
than the others, and measures 36 mm. in length, whereas the pos* 
terior petals of the trivium measure 28 mm. in length. The petals 
of the bivium, or I and V, are longer than the posterior pair of the 




v; 



Fig. 2. — Encopb platttata. Dhawixq of the typb-specimen, natural size. Restora- 
tions ARE INDICATED BY DOTTED LINES. 

trivium, but as they are incomplete posteriorly, a measurement can 
not be given. On the ventral side, the ambulacral furrows are 
strongly marked and each gives off a few weakly impressed branches. 
The apical disk is quite well preserved, shows clearly the ocular 
pores and four of »the five genital pores, which are a characteristic 
feature of Encope. The only genital pore wanting is that occurring 
in area 1, which is destroyed by a local fracture of the test. Minute 
tubercles cover the dorsal side of the test. Ventrally the tubercles 
are larger except near the ambulacral furrows where they are minute. 



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110 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Encope platytdta is a near ally of Encope tenuis Kew* of the 
Miocene of California, but differs from that species in that the great- 
est height of the test is central, and the periproct is confluent with 
the lunule. 

Locality and geologic occurrence. — Gatun formation, Miocene, 
Panama Canal Zone, from lowest horizon in big cut, one-fourth to 
one-half mile beyond Camp Cotton toward Monte Lirio, D. F. Mac- 
Donald and T. W. Vaughan, collectors, 1911, U. S. National Museum 
station No. 6029a, one specimen. 

Type.— Cfki. No. 324455, U.S.N.M. 

ENCOPE HEGATREMA JackMii. 

Plate 52, fig. 1. 
Encope megatrema Jackson, Proc. U. S. Nat Mus., vol. 53, 1917, p. 496, 
text figs. 3, 4 ; pi. 68, fig. 1. 

This species is represented by one fairly good test with its counter- 
part, and in addition some 12 fragments which yield helpful facts 
on close study. From the incompleteness, measurements and some 
details will have to be given in general terms or omitted. As a whole, 
the test is low, elongated, thin on the borders and with shallow 
ambulacral notches and an enormous lunule in interambulacrum 5. 

From the best specimen, which is figured, the length probably was 
about 120 mm. and the width about 106 nmi. ; thickness of the test 
at its center is 10 mm. Ambulacral notches are shallow and quite 
wide in areas II and V, indicating that this is the character in the 
two posterior ambulacra I and V and also in the paired anterior am- 
bulacra II and IV. This evidence is supported by several of the frag- 
ments which show shallow lobes like the type, but it can not be 
definitely stated which areas they represent. The notch of the ante- 
rior odd ambulacrum III is not known, but it was probably shallower 
than the others, as is characteristic of species of the genus. The most 
striking feature of this species is the lunule in interambulacrum 5, 
which is enormous. It is situated about midway between the apical 
disk and posterior limits of the test, and is roughly triangular in 
shape, the apex of the triangle pointing anteriorly. It measures at 
the surface of the opening 27 mm. in length and 27 mm. in width at 
the widest part posteriorly. The walls of the lunule slope outward 
from the center, as seen looking from above, as is well shown in two 
of the fragmentary specimens. From this sloping character of the 
walls, it results that the width of the lunule would be greater by about 
6 to 10 millimeters on the ventral side than it is on the dorsal. The 
height of the wall of the lunule is 12 mm., which is doubtless the 
highest point of the test. The lunule in this species is, relatively to 
the size of the specimens, the largest known in any species of the 

» Kew. W. S. W. Tertiary echlnolds of the Carrlzo Creek Region In the Colorado Desert. 
University of California Bull., Dept. Geology, vol. 8, No. 5, pp. 39-60, pis. 1-S, 1914. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 



Ill 



genus, fossil or living. It is striking that this great size of the lunule, 
a progressive character, should be associated with small and shallow 
ambulacral notches which, for the genus, is a relatively primitive 
character. 

The ambulacral petals are beautifully distinct and well preserved 
for part of their extent in the type and one other specimen. The 




-^ ^ ^ 



Fig. 3. — a, Bncopb megatuema. TToi-otype, natural size. The akea showing part of 

AMBULACRUM II IS DRAWN FROM A rUAC.MENT. RESTORATIONS ARE INDICATED BY DOTTED 

LINKS, ft. Section of luxule to show the inclined faces, drawn from two 

FRAGMENTS. 

posterior pair, I and Y, are long and narrow with a relatively wide 
poriferous area and narrow median interporiferous area. The width 
of the petal of ambulacrum V is 11 mm. and its length is 50 mm. It 
extends posteriorly in a wide curve around the lunule of interambu- 
hicrum 5 and reaches a line coincident with the posterior end of the 



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112 BULLETIN 103, XJXITED STATES NATIONAL MUSEUM. 

lunule. It also extends to within 5.5 of the ambulacral notch of the 
area in which it lies. Ambulacrum IV is much shorter than ambula- 
crum V, measuring 30 mm. in length and 12 mm. in width at its 
widest part. This ambulacrum extends to within 3.5 mm. of its 
marginal notch. The features of the ambulacra V and IV, as de- 
scribed, indicate the character of ambulacra I and II, which are only 
preserved in part in the holotype, though one of the fragments has 
ambulacrum II quite perfectly presened. Ambulacrum III is repre- 
sented only in part (for a length of 25 mm.) by the left side of its 
petaloid area; it probably had about the length and width of the 
petal of ambulacrum IV, as in the jiUied species Encope macrophora 
Ravenel. Ambulacral furrows on the ventral side are deep, strongly 
marked, with some forking near the periphery of the test. The inter- 
ambulacra are very wide, not narrowing markedly near the apical 
disk. Minute tubercles cover the dorsal surface of the test, and ven- 
trally the tubercles are larger excepting on the lines of ambulacral 
furrows, where they are minute or wanting. Details of the apical 
disk, peristome and periproct are entirely wanting. This species does 
not make a close approach to any other known species, but its nearest 
ally is Encope macrophora Eavenel from the upper Miocene of 
South Carolina and the Pliocene of Florida.^ 

Locality arid geologic occurrence, — Gatun formation, Miocene, 
Panama Canal Zone. From 85-foot cut north side of big swamp on 
relocated line, Panama R. E., about one and one-half to two miles 
beyond Camp Cotton, toward Monte Lirio, D. F. MacDonald and 
T. W. Vaughan, collectors, 1911. Fourteen specimens, including 
fragments, U. S. National Musemn station No. 6030. 

Type.— Cut No. 324456, U.S.N.M. 

ECHINOLAMPAS SEMIORBIS Gupp^. 

Echinolampaa scmiorbis Guppy, On Tertiary Echlnoderms from the West 
Indies, Quart. Journ. Geol. Soc. London, vol. 22, 1866, p. 299, pi. 19, 
fig. 7. — CoTTEAu, Ecliinides Tertiares des lies St Barth^lemy et An- 
guilla, Kongl. Svenslv. Vetcnskaps. Akad, vol. 13, 1875, p. 24, pi. 5. 
figs. 1-2; pi. 6, fig. 1.— -Jackson, Proc. U. S. Nat. Mus., vol. 53, 1917. 
p. 498. 

This species is abundant in the Oligocene Tertiary of the West 
Indies, material from Anguilla having been described by Guppy, and 
Cotteau erroneously records it from St. Bartholomew. Dr. T. Way- 
land Vaughan in 1914 collected abundant, fine specimens in the Island 
of Anguilla. 

From the Panama Canal Zone a number of specimens were col- 
lected from a hard gray limestone. The specimens are for the most 
part uncompressed and in very good condition of preservation. One 



1 Clark, William Bullock, and Twltchell, Mayvlllc Vf. Mesozolc and Cenozolc Echino- 
dermata of the United States. Monograph, U. S. Geol. Survey, vol. 64, 1915, p. 206, 
pi. 03, flga. 2a-e; pi. 04, figs, la-f. 



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GEOLOGY AND PALBOKTOLOGY OF THE OAKAL ZONE. 118 

of the largest specimens measures 107 mm. in length, 103 mm. in 
width, and 58 mm. in height 

Locality and geologic oocurrenoe. — ^Upper Oligocene. Emperador 
limestone, Panama Canal Zone. Upper Limestone bed near Tower 
*'N" (opposite Las Cascadas, Gaillard cut), D. F. MacDonald, col- 
lector, 1911, U. S. National Museum station No. 5866&, one specimen, 
U. S. Nat. Mus. Cat. No. 324457. Also Panama Canal Zone, from 5th 
or topmost limestone, Gaillard cut, opposite Las Cascadas, U. S. 
National Museum station No. 6019gr, D. F. MacDonald and T. W. 
Vaughan, collectors, 1911, 4 specimens. U. S. Nat. Mus. Cat. No. 
324458. 

SCHIZASTER ABMIGER W. B. CUrk. 

Schiz<Mter armiger Clabk and Twitchell, Mesozoic and Oenozoic Echlno- 
dermata of the United States, Monograph U. S. Geol. Survey, vol. 64, 
1915, p. 152, pi. 70, figs, la-d.— Jackson, Proc. U. S. Nat Mus., vol. 
53, 1917, p. 498. 

In this species the test is rather large, cordiform; upper surface 
slopes at first rapidly, then more slowly from the anterior margin to 
the apical system beyond which an elevated sharp ridge continues to 
the truncated posterior margin. Length, 59 mm. ; width, 50 mm. ; 
height, 25 mm. The ambulacra are broad and the odd anterior 
ambulacral petal III is situated in a deep groove that indents the 
anterior margin. The two lateral anterior ambulacra II and IV are 
in deep, broad grooves, with petals 18 mm. long. The posterior 
ambulacra I and V, similar but shorter, are 9 mm. long. Peripetalous 
fasciole is broad and distinct. Interambulacra gibbous, the posterior 
No. 5 being built up into an elevated keel. The peristome is indis- 
tinct in our specimen, but as shown in W. B. Clark's excellent figures, 
is wide and near the anterior margin. The periproct is high on the 
truncated posterior end. 

The type material described by Clark is ascribed to the upper 
(Jackson) Eocene of Choctaw County, Alabama. 

Locality and geologic occwrrence, — ^Miocene (?),^ Bonilla, Costa 
Rica, Hill collection, U. S. Nat. Mus. Cat. No. 135214, one specimen. 

SCmZASTER CRISTATUS JackMii. 

Plate 52, figs. 2-4. 

Schizaster cristatus Jackson, Proc. U. S. Nat. Mus., vol. 53, 1917, p. 499, 
pi. 68, figs. 2-4. 

The material of this species consists of two internal moulds ; as the 
plates are entirely wanting, of course external characters can not be 
given. The more perfect of the two specimens measures 40 mm. in 
length, 36 mm. in width, and 22 mm. in height Test is moderate 

^ According to Hill and DaU the rocks exposed at this locality are of the same age as 
those at Oatan, Canal Zone. FOr a further discussion see the last chapter (by Vaughan) 
in Uiis Tolume. 



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114 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

sized, cordifann, sloping gradually from the anterior border up to the 
median crest, the widest portion being through the middle of the test. 
The most striking feature of this species is the median keel-like crest 
that rises sharply from the summit of interambulacrum 5 at the pos- 
terior border of the test. 

The petal of ambulacrum III is sunken in a deep, wide groove, ex- 
tending to the anterior border of the test and measuring 23 mm. in 
length. The petals of the lateral anterior ambulacra II and IV are 
in deep grooves measuring 18 mm. in length and having about 22 
plates in. each half ambulacrum, as is indicated by the casts of the 
pores. The petals of the posterior ambulacra I and V are widely 
divergent from the anterior pair, nearly parallel and directed back- 
ward in deep, sunken grooves. The grooves are 7 mm. long, and 
there are about 14 plates in each half ambulacrum at this point, as 
indicated by casts of the pores. The periproct is situated on the 
posterior face and coincides with the base of the crest in interam- 
bulacrum 5. The peristome is wide and situated far forward, the 
tip which abnost closes the mouth being 10 mm. from the anterior 
border of the test. 

Locality and geologic occurrence. — Miocene( ?), Brazil, Costa Rica. 
A. Alfaro, collector, U. S. National Museum station No. 5505, two 
specimens. 

Type.— C^t. No. 324460, U.S.N.M. 

SCmZASTEB PANAMEN8I8 Jacluon. 

Plate 50, ftjrs. 2-3. 

Schizaster panamensis Jackson, Proc. U. S. Nat. Mus., vol. 53, 1917, p. 500, 
pi. 66, figs. 2, 8. 
The material consists of an internal mould free from matrix, and 
three specimens more or less Complete, embedded in porous, dark- 
colored volcanic tuff which also bears some fragments of lignite. 
The specimen, free from matrix, is the most completely preserved, 
although somewhat compressed dorso-ventrally, and is selected as 
the type. The specimen measures 48 mm. in length, 40 mm. in width, 
and 25 mm. in height The petals of the ambulacra are situated in 
broad, deep furrows. The anterior petal III extends to the anterior 
limit of the test and measures 23 mm. in length. The paired anterior 
ambulacra II and IV are in grooves 13 mm. long and diverge widely 
from the anterior petal. The posterior petals I and V are shorter 
than the anterior pair, measuring 5 mm. in length, and are directed 
almost straight backward. The position of the periproct is not 
clearly indicated on the mould, but apparently it is near the upper 
part of the posterior face. The peristome is wide and rounded, and 
is situated 16 mm. from the anterior border of the test. The peri- 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 115 

petalous fasciole is quite wide and is fairly well shown in areas 11 
and rV on the type-specinien and still better in one of the fragments, 
which is a counterpart of the dorsal side of the same. 

One of the specimens, which is an eicternal mould, shows the im- 
press of the outline of the plates of part of a test, and gives measure- 
ments of considerable interest. It measures about 50 mm. in length, 
about 46 nun. in width, and about 38 mm. in height. From incom- 
pleteness of the specimen no exact measurements can be given, yet 
those available indicate a very high test. 

Locality and geologic occurrence. — Gatun formation, Miocene, 
Panama Canal 2k>ne. Second cut, south of Oatun R. R. Station, 
Goldman Coll., U. S. National Museum station No. 7294, four speci- 
mens. Holotype, U. S. Nat. Mus. Cat. No. 324461. Another speci- 
men, imperfect and much worn, with a very high test, and appar- 
ently referable to this species, is from Panama Canal Zone, Gatun, 
section A, from middle of Bed " E,'' D. F. MacDonald, collector, U. S. 
National Museum station No. 6008, one specimen. U. S. Nat Mus. 
Cat. No. 324462. 

DESCRIPTION OF PLATES. 

Plate 46. 

Pia 1. Clypeaster lanceolatua Otteau, dorsal view, natural size. The dark 
spot in interambulacram 5 is a yeUow label that took black in the 
photograph. : U. S. Nat Mus., Oat. No. 824451, Station 58665. 
2. Another specimen of the same, ventral view, natural size, U. S. Nat 
Mus. Cat. No. 324451, Station 58666. 

Plate 47. 

Fig. 1. Clypeaster ffatuni Jackson, dorsal view. Holotype, slightly reduced^ 
U. S. Nat Mus. Cat No. 324453, Station 5662. 

Plate 48. 

PiQ. l« Olypeaster gatwii Jackson, ventral view; same specimen as Plate 47. 
Holotype, slightly reduced, U. S. Nat Mus. Cat No. 824452, Station 
5662. 

Plate 49. 

Fio. 1. Enoope annectans Jackson, dorsal view, natural size. Holotype, U. S. 
Nat Mus. Cat No. 324454, Station 5846. 
2. The same, ventral view. 

Plate 50. 

Fro. 1. Enoope annectana Jackson, another specimen, external mould of ven- 
tral side seen from above. Natural size, Paratype, U. S. Nat Mus. 
Cat No. 824466, Station 5846. 

2. ScMzoBter panamenaia Jackson, dorsal view, natural size. Holotype, 
U. S. Nat Mus. Cat No. 324461, Station 7294. 

8. The same, ventral view. The dark spot in interambulacram 5 is a yel- 
low ticket that took black in the photograph. 



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116 BULLETIN 103, TJNITBD STATES NATIONAL MUSEUM. 

Plate 51. 

Fig. 1. Encope platytata Jackson, dorsal view, natural size. Holotype, U. S 
Nat Mus. Oat No. 824455, Station 6029a. 
2. The same, ventral view. The dark spot In Interambulacmm 4 of fis. 2 
and in interambulacrum 2 of fig. 2 are yellow tickets that took bla<A 
in the photographs. 

Plate 52. 

Fig. L Encope megatrema Jackson, dorsal view, natural size. Holotype, natu: 
ral size, U. S. Nat Mus. Cat No. 324456, Station 6030. 

2. Sohizaster cristatus Jackson, dorsal view, natural size. Holotype 
U. S. Nat Mus. Cat. No. 324460. Station 5505. 

8. The same, ventral view. 

4. The same, side view. 



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U. 8. NATIONAL MUSEUM 



BULLETIN 103 PL. 46 




CM = 

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O 
LU 
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-^ < 

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U. 8. NATIONAL MUSEUM 



BULLETIN 103 PL. 47 




Dorsal View of Clypeaster qatuni. 

For explanation of plate see paoe 1 1 5* 



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U. S. NATIONAL MUSEUM 



BULLETIN 103 PL. 48 





^m^^^ 



Ventral View of Clypeaster qatuni. 

For explanation of plate see paqe 116. 



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U. & NATIONAL MUSEUM 



BULLETIN 103 PL. 49 




Dorsal (I) and Ventral Views (2) of Encope annectans. 

Fon EXPLANATION OF PLAT! •■■ PAOB 116. 



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U. S. NATIONAL MUSEUM 



BULLETIN 103 PL. 60 






Mould of Ventral Side of Encope annectans seen from above. (2) Dorsal, 
AND (3) Ventral Views of Schizaster panamensis. 

For explanation of plate see paqi us. 



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U. 8. NATIONAL MUSEUM 



BULLETIN 103 PL. 61 




) 



/ 



^■^ . 




Dorsal View (I) and Ventral View (2) of Encope platytata. 

For explanation of plate see paqe 116. 



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U. S. NATIONAL MUSEUM 



BULLETIN 103 I PL. 62 






4 



I. Dorsal View of Encope megatrema. Dorsal (2), Ventral (3), and Side (4) 
Views of Schizaster cristatus. 

For explanation of plate sec page lie 



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INDEX. 



Page. 

annectans, Bncope 108,106 

Annlger, Schlxaster 104, 118 

bowcni, Clypeaster 106 

ClTpeaster bowersi 105 

gatanl 108. 105 

lanceolatns 108, 104 

criatatni, Schlzaster 104,118 

EchlnolampaB semlorbis 104, 112 

Encope annectans 108,106 

grandis 108 

macrophora 112 

megatrezna 104, 110 

mlchellnJ 108 

micropora 108 

pUtytato 108, 108 

tenulB 110 



Pagtu 
gatiml, Clypeaster 108,105 

grandis, Bncope 108 

lanceolatns, Clypeaster 108,104 

macrophora, Bncope 112 

megatrema, Bncope 104, 110 

Mellita sezlesperforata • 107 

michellnl, Bncope 108 

micropora, Bncope 108 

panamensis, Schlsaster 104,114 

platytata, Bncope 108, 108 

Schlsaster armlger 104, 118 

crUtatus 104, 118 

panamensis 104,114 

semiorbis, Bchlnolampas 104, 112 

sezlesperforata, Mellita 107 

tennis, Bncope 110 

X 



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SMITHSONIAN DJSTITtrnON 

UNITED STATES NATIONAL MUSEUM 

Bulletin 103 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



BRYOZOA OF THE CANAL ZONE 
AND RELATED AREAS 

By FERDINAND CANU 
Of VersailUs, France 

AND 

RAY S. BASSLER 
Of Washington, District of Columbia 



Extract from Bulletin 103, paget 117-122, widi Plate S3 




WASHINGTON 

COVERNMtNT PRINTING OFHCE 

1918 



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BRYOZOA OF THE CANAL ZONE AND RELATED 

AREAS. 



By Ferdinand Canu 

0/ Versailles, France 

AND 

Ray S. Basslbr 
Of Washington, D^trict of Columbia, 



The following pages contain the descriptions of the few bryozoa 
that have so far b^n found in the rocks of the Canal 2ione and 
related areas. These bryozoa consist of two species from the £m- 
perador limestone of the Canal Zone collected by Messrs. T. Way- 
land Vaughan and D. F. MacDonald and three species from the 
Miocene of Costa Rica collected by D. F. MacDonald. The list of 
species described is as follows : 

Ogivdma rrmtabUis^ new species, Emperador limestone, Panama 
Canal Zone. 

Holoparella alhirostris (Smitt), Emperador limestone, Panama 
Canal Zone. 

Cupularia vmbeUata Defrance, Miocene, Costa Rica. 

Cuptdaria canariensis Busk, Miocene, Costa Rica. 

Stichoporina tuherom^ new species, Miocene, Costa Rica. 

Order CHEILOSTOMATA. 

Group MEMBRANIPORAE. 

Genoa OGIVALINA Canu and BaMler. 

OGIVALINA MUTABIUS. mw spedet. 

Plate 58, fig. 1. 

The zoariimi is incrusting. The zooBcia are elongated, oval, dis- 
tinct, separated by a deep furrow; the mural rim is thin, smooth, 
rounded; there is often a small gymnocyst. The opesium is very 
large, irregular, more often oval. The ovicell is endozooecial, small, 

S370d— 18 117 



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118 BULLETIN 103; UNITED STATES NATIONAL MUSEUM. 



Measurements. — Opesium 
Zooecium 



little convex. Sometimes there is a small interzooBcial fusifoim 
avicularium (?). 

fA^=0.6(M).70mm 

/^=0.30-0.45 mm 

Z;3=0.75-0.95mm 

fe=0.5(M).70mm 

The *great irregularity of form and zooecial dimensions of this 
species justifies its name. There are some variations which recall 
those of Membranipora irregularis Manzoni, 1875 ^ which possesses 
a mural rim enlarged at the base and also some large interzooeciai 
avicularia. 

The present species differs from the splendid Ogivalina eximipora 
Canu and Bassler from the Middle Jacksonian of North and South 
Carolina in its smaller dimensions, in the absence of cryptocyst, and 
in the presence of a gymnocyst. The avicularium (?) is identical 
in form and position. 

Occurrence. — ^Emperador limestone, old quarry, one-third mile 
north of west of Empire, Panama Canal Zone. D. F. MacDonald 
and T. Wayland Vaughan, collectors, 1914, Station No. 6016. 

Southwest side Crocus Bay Hill, Anguilla, Leeward Islands. 
T. Wayland Vaughan, collector, 1914, Loc. No. 6893. 

Holotype.—CdX. No. 65039, U.S.N.M. 

Family OPESIULIDAE Jullien. 
Geniifl CUPULARIA Lamonroiix. 

GUPULARIA UMBELUITA Defnnce, 181S. 

Plate 53, figs. 2-4. 

1908. Cupularia umhellata Canu, Iconographie des Bryozoaires fossiles 
de TArgentiDe, Anales del Museo Nacional de Buenos Aires, vol. 17, 
p. 275, pi. 5, figs. 4, 5. (See for complete bibliography.) 

1909. Cupularia umhellata Canu, Bryozoaires fossiles du Sad-Ouest de 
la France, BaUetin de la Soci^t^ Geologique de France ser. 4, vol. 9, 

. pp. 448, 464, pi. 16, figs. 16, 17. (Regional bibliography.) 
1909. Cupularia lotoei Norman, On the Polyzoa of Madeira, Journ. Lin- 

nean Soc, vol. 80, p. 290, pi. 87, figs. 7-12. 
1918. Cupularia umhellata Canu, £tude co;npar^ des Bryozoaires Hel- 
vetiens de I'figypte avec les Bryozoaires vivants de la Mediterran6e 
et de la Mer Rouge, Mem. a I'instltut Egyptien, vol. 6, fasc. 3, p. 205. 

1913. Cupularia umhellata Canu, Cont. a Tetude des Bryozoaires fossiles, 
pt 5, p. 126; pt 7, p. 126; pt 12, p. 127; Bulletin Soc. Qeoh France 
(IV, XIII). 

1914. Cupularia lowei Osbubn, The Bryozoa of the Tortugas Islands, 
Florida. Publication No. 182, of the Carnegie Institution of Washing- 
ton, p. 194. 

» BrioEoi del pliocene antico dl Castrocaro, Bologna, 1876, p. 10, pi. 1, flge. B, 8. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 119 

The fossils which are identified as above are rather well preserved 
and their determination is easy. The pores of the hydrostatic zooecia 
are not radicular. We are ignorant as to why Norman, who is a 
great lover of archaic names, has not preserved the name of Def ranee. 
The figures published by this author and by d'Orbigny are excellent 
and leave no doubt as to the identity of the two species. 

Occurrence. — ^Miocene, Banana River, Costa Bica. D. F. Mac- 
Donald, collector, 1911. Bowden marl, Bowden, Jamaica. 

This species is almost always associated with C. canariensia Busk. 
Like the latter, it commences in the Alum Bluff formation and con- 
tinues in the higher Miocene and Pliocene deposits of the United 
States. 

Oeological distribution. — ^Aquitanian of Italy (Seguenza, Nevi- 
ani), of Bordeaux (Canu). Burdigalian of Italy (Seguenza, Canu), 
of Bordeaux (Canu). Helvetian of Italy (Seguenza), of Touraine 
(Canu), of Bordeaux (Canu), of Maryland (Ulrich), of Egjrpt 
(C!anu). Tortonian of Provence (Canu), of Italy (Seguenza). 
Plaisancian of England (Busk), of Italy (Manzoni). Astian of 
Italy (Neviani, Canu), of Provence (Canu). Sicilian of Italy 
(Neviani). Quaternary of Italy (Seguenza), of England (Canu). 

Habitat. — ^Mediterranean. Atlantic to the Canary Islands, and 
Florida. It is common in the Gulf of Gascony in the Miocene; it 
has now disappeared from it. 

It has been dredged at a depth of 11 to 48 meters in America and 
from 81 to 118 meters in Madeira. 

CUPULASIA CANABIEN8I8 Buk. 

Plate 53, figs. 5-7. 

1906. Cupularia oanariensU Canu, Iconographie des Bryozoaires fossilea 
de TArgentlne, Anales del Musee Nacional de Buenos Aires, vol. 17 
(aer. 8, vol. 10), pt. 1, p. 275, pi. 5, figs. 8, 9, 10. (See for complete 
bibliography.) 

1900. Cupularia guineensis Nobman, The Polyzoa of Madeira and neighbor- 
ing islands, Linnean Society's Journal, Zool., vol. 80 (July), p. 289, 
pi. 37, figs. 2rS, 

1914. Cupularia guineensis Osbubn, The Bryozoa of the Tortngas Islands, 
Florida, Publication No. 182, of the Carnegie Institute of Washington, 
p. 194. 

The beautiful figure published by Busk in 1859, has led all paleon- 
tologists to use the specific term ccmariensiSy especially since the same 
iQthor distinguished thia species from Ouptdaria guineenais Busk, 
1854. For a half century, it was therefore employed by Busk, Waters, 
Kanzoni, Van den Brock, Neviani, Seguenza, De Angelis, and Canu. 

Kow it appears established that Busk's two species are identical 
(Norman, Osbum). We do not believe it necessary to change the 



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120 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

names of these species as the latter authors have done since the 
author of each is the same. A simple question of date ou^t not 
alter all the literature of this species which although it has never 
been entirely published is nevertheless quite important. 

Our American specimens are well preserved. 

Occurrence, — ^Miocene, Banana River, Costa Rica. D. F, MacDon- 
aid, collector, 1911. Bowden marl at Bowden, Jamaica. 

The earliest occurrence of this species in the United States is in the 
Alum Bluff formation, but it is found also at many other horizons of 
the Miocene and Pliocene. 

Geological distribution. — Burdigalian of Bordeaux (Collection 
Canu). Helvetian of France (Canu) of Spain (De Angelis). Tor- 
tonian of Austria-Hungary (Reuss), of Italy (Seguenza). Plais- 
ancian of Italy (Manzoni), of England (Busk), of Spain (De An- 
gelis), of Algeria (Canu). Astian of Italy (Neviani, Canu). Sicil- 
ian of Rhodes (Manzoni), of Italy (Neviani). Quaternary of Italy 
(Neviani), of Argentina (Canu). Miocene of Australia? (Waters). 

Family CELLEPORIDAE Busk. 
Geniifl HOLOPORELLA Waters. 

HOLOPORELLA ALBIROSTRIS (Smitt). 

Plate 53, fig. 8. 

1873. Discopora albirostris Smitt» Floridan Bryozou, pt. 2, Kongl. Svenska 

Vetenskaps-Akademlens Handllnjrjir, vol. 11, No. 4, p. 70, pi. 12, figs. 

233-239. 
1889. Cellepora alhirostrift Jelly, A Sjoionymic Catalogue of the Uecent 

Marine Bryozoa, p. 45. (See for complete bibliography.) 
1914. HoloporeUa alhirostHn Osbubn, Bryozoa of Tortugas Islamls, Pub. 

182. Carnegie Institution, p. 215. 

Of the two Specimens of this species which have been collected at 
Panama and at Anguilla one corresponds to Smitt's figure 237 and 
the other to figure 238. 

Occurrenx:e, — ^Rare in the Emperador limestone at the old quarry 
one-third mile north of west of Empire, Panama Canal Zone, D. F. 
MacDonald and T. Wayland Vaughan, collectors, 1911 (Station 
No. 6106). Also rare along the southwest side of Crocus Bay, An- 
guilla, Leeward Islands, Dr. T. Wayland Vaughan, collector^ 1914, 
Loc. No. 6894. 

Geological distribution. — Miocene of Australia and New Zealand 
(Waters). Habitat. Atlantic off Florida. Pacific off Australia. 
Specimens have been dredged off Australia to a depth of 121 meters. 
Smitt in Florida has discovered them between 40 and 56 meters, but 
Osbum states that it abounds at a depth of 24 meters. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 121 

Family CONESCHARELLINIDAE Levinsen. 
Genufl STICHOPORINA StoUczka. 

STICHOPORINA TUBEROSA. new speciM. 
^ Plate 53, flgs. 9-12. 

The zoarium is free, conical, hollow with very thick walls. The 

peristome is salient, ornamented with small tuberosities; it bears one 

or two small elliptical avicularia with bar or denticles. The ovicell 

is large, somewhat salient, convex; it is hyperstomial and always 

closed by the operculimi. On the lower face, there are large pores 

surrounded by very small ones. 

-MM a A _x f A«=0.15 mm. 

Measurefnents, — Apertura K . ^ nn 

^ |7^=0.09 mm. 

This is a very elegant species characterized by its peristomial tuber- 
osities. The ancestrula is visible only in the interior of the zoarium ; 
it is covered exteriorly by the first zooecia. All the zooecia are sepa- 
rated from each other by small canals which appear to end in the 
large, inferior pores. 

This species must not be confounded with Mamillopora cupula 
Smitt, 1872. It diflfers from it in its ovicell which is not bilobate 
and in its ovarian zooecia which are not larger than the others. 

Occurrence, — Miocene, Banana River, Costa Rica, D. F. McDonald, 
collector, 1911. 

Cotypes.—Cfit No. 65040, U.S.N.M. 

EXPLANATION OF PLATE 53. 

OgiraHna mutabilis, new species. 

Fio. 1. The type-specimen, X 20, with large Irregular opesia, small ovicell, small 
gymnocyst and one zcecium with a fusiform aviciilarium. 
Eroperador limestone. Crocus Bay Hill, Anguilla. 

Cupularia nmbellata Def ranee. 

Fig. 2. Two zoaria, natural size. 

3. CeUuliferous convex surface, X 20. 

4. Concave surface, X 20. 
Miocene, Banana River, Costa Rica. 

Cupularia canariensis Busk. 

Fio. 5. Two zoaria, natural size. 

6. CeUuliferous convex surface, X 20. 

7. Concave surface, X 20. 
Miocene, Banana River, Costa Rica. 



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122 BULLETIK 103, UNITED STATES NATIONAL MUSEUM. 

Holoporella albirostris (Smitt). 

Fig. 8. Several soceda much enlarged (after Smitt). 
Recent, Gulf of Mexico. 

SUchoporina tuberosa, new species. 

Fig. 9. Two zoaria, natural size. 

10, 11. Two views X 20, of the convex, ceUullferous side. 
12. Photograph of the concave side, X 20. 
Miocene, Banana River, Costa Rica. 



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U. S. NATIONAL MUSEUM 



BULLETIN 103 PL. 53' 




Bryozoa of the Panama Canal Zone and Related Areas. 

For exFLANATiON of platc see PAQES 121. 122. 



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INDEX. 

f Synonyms are in Italics.] 



Page. 
•JUrotrls, Cellepora 120 

DUoopora 120 

tIfalroctrU, HoloporeUa 117,120,122 

eanariensto, Cnpularia 117, 110, 121 

CeOepara ailfiro9tri9 120 

Cnpnlaria c&narlensU 117, 110, 121 

Cnpuktria gwtne&mHe 110 

lotoei 118 

Cnpulmria nmbellata 117,118,121 



Pa£9. 

Difioopora allAroMtriB 120 

guineen»i9, Oupularia 110 

HoloporeUa albirostris 117,120,122 

loioei, Oupularia 118 

mutabills, Oglvalina 117, 121 

OglyaUna mutabillB 117, 121 

Stichoporlna tnberosa 117, 121, 122 

tuberosa, Stichoporlna 117, 121, 122 

nmbellata, Cnpnlarla 117, 118, 121 

I 



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SMITHSOmAM iNsTrrvnoM 

UNITED STATES NATIONAL MUSEUM 

Bulletin 103 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 






I 



DECAPOD CRUSTACEANS FROM THE 
PANAMA REGION 



By MARY J. RATHBUN 
Associate in Zoology, United States National Museum 



Extract (rom Bulletin 103, pages 123-184, with Plates 54-6 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1918 



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I 

I 



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SMITHSONMN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 
Bulletin 103 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



DECAPOD CRUSTACEANS FROM THE 
PANAMA REGION 



By MARY J. RATHBUN 
Associate in Zoology, United States National Museum 



Eitract from Bulletin 103, pages 123-184, with Plates 54-66 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1918 



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DECAPOD CEUSTACEANS FROM THE PANAMA 

REGIOK 



By Mabt J. Rathbun, 
Atiociate 4n Zoology f United States National Museum. 



INTRODUOTION. 

Fifty-eight species of Decapods are enumerated from the collec- 
tions examined by the author. Three species described by other 
authors are inserted in systematic order, thus making the list com- 
plete to date for the Panama region. All the available material 
in the United States National Museum from Panama and Costa Bica 
is included; it ranges in age from the Oligocene (Culebra formation) 
to the Pleistocene. 

In the list of stations and the table of distribution the data relat- 
ing to Cirripedia from Dr. H. A. Pilsbry's report are included for 
convenience of reference. 

The literature on Panama Tertiary Decapods is so scanty that it 
IB not surprising that nearly all of the forms now examined prove to 
be new. Six species previously described from living forms are here 
recorded from the Pleistocene (4 species) or the Pliocene (2 spe- 
cies). Thirty-nine species are described as new, three are types of 
new genera, and one of these is the type of a new family, the Ght- 
tnniidae. This is an extremely large and massive crab and combines 
the characters of the well-known Recent families, the Cancridae and 
the Portunidae. The most remarkable occurrence is that of a mem- 
ber of the Hexapodinae, that subfamily of the Ooneplacidae in which 
the legs of the last pair are wanting. This is a small group of Recent 
crabs containing 5 genera and 8 species and is strictly Indo-Pacific. 
The species from the Oligocene of Panama is the first one observed 
in a fossil state. Many other genera dealt with in this report have 
never before been found fossil. Such are PachyrhdeSy PetroUsthes^ 
Axiusj HepatuSy Mursia^ LeucosUia, Euphylam^ Heteractaea^ Ev/ry- 
^tMw, EuryplaXy and Cardiaoma. 

As in all large collections of fossil crustaceans there are a number 
of fragments whose position is problematic Some of these can be 
determined as to genera, others as to. family only. 

83706— 18— BuU. 108 128 



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124 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

LITERATURE OK TERTIARY DECAPODS OP PANAMA. 

BouviBR, E. L. Calappa Zurcheri, Crabe nouveau des terrains mio- 
cfenes de Panama. Bull. Mus. Hist Nat. Paris, vol. 5, 1899, pp. 
189-192, 1 text-fig. 

Calappa zurcheri is not represented in tlie United States Geological Survey 
collections. 

TouLA, Franz. Die jungtertiare Fauna von Gatun am Panama- 
kanal. II. Teil. Jahrbuch der k. k. Geolog. Reichsanstalt^ 
Wien, vol. 61, 1911, pp. 487-530 (1-44), pis. 30, 31 (1, 2). 

The hermit-crab {Petrochirus) noted and figured by Toula (p. 511, pi. 30, fig. 
13) I have ventured to describe as a new species, combining as it does the char- 
acters of the two nearly related Recent species which inhabit opposite sides of 
the continent The ** Krabbenscheren " of Toula (p. 512, pi. 30, fig. 14) are de- 
scribed below as a species of Callianassa, C, toulai, 

Bbown, Amos P., and Pilsbry, Henry A. Fauna of the Gatun For- 
mation, Isthmus of Panama. II. Proc. Acad. Nat. Sci. Phila- 
delphia, vol. 64, Dec. 1912 (publ. Jan. 30, 1913), pp. 500-519, pis. 
22-26. 

The author is indebted to Dr. H. A. Pilsbry for the loan of the specimens of 
CMianasaa in the collection of the Philadelphia Academy of Natural Sciences 
which were described by Brown and Pilsbry. They have been critically com- 
pared with those collected by the United States Geological Survey. 

LIST OF STATIONS FROM WHICH MATERIAL HAS BEEN EXAMINED, 
ARRANGED FROM THE EARLIEST TO THE LATEST, WITH THE 
SPECIES FOUND AT EACH. 

Station 601Sa^ — Panama Canal Zone. One-quarter mile south of 
Empire Bridge. From lower dark clay beneath lower conglomerate. 
Culebra formation (lower part). Oligocene series. Collectors^ 
D. F. MacDonald and T. W. Vaughan; 1911. Balaniis {Hesperi- 
halanus?)^ species. Callianassa lacunosa Rathbun. 

Station 6010. — Panama Canal Zone. Near Panama Canal Station 
"1910," north of Pedro Miguel locks. From dark clay. Culebra 
formation (lower part). Oligocene series. Collectors, D. F. Mac- 
Donald and T. W. Vaughan; 1911. Mvrsia ohsciira Rathbun. 

Specimens in Musewm^ Academy of Natural Sciences^ Philadel— 
pMa. — ^Panama Canal Zone. Las Cascadas section, Gaillard Cut. 
Lignitic layers about 65 feet below the base of Pecten bed at Tower 
N. Culebra formation (central part). Oligocene series. Collector, 
Prof. William B. Scott; 1911. Callianassa scotti Brown and Pils- 
bry. CaJUanassa spinulosa Rathbun. Callianassa guadrata Rath- 
bun. 

1 The station numbers refer to the station book of Cenozolc Inyertebrate fossils of the 
United States National Museum. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 125 

Station 6019b. — ^Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. Four feet of dark, stratified tuff and clay immediately 
overlying the lower limestone bed. Culebra formation (upper part). 
Oligocene series. Collectors, D. F. MacDonald and T. W. Vaughan; 
1911. CaUianassa scotti Brown and Pilsbry. Callinectes^ species, 
Panopew, species. 

Specimen in Musewrn^ Academy of Natural Sciences^ Philadel- 
phia. — Costa Bica. Probably Culebra formation. Oligocene series; 
labeled "Miocene." Collector, W. M. Gabb. CaUianassa scotti 
Brown and Pilsbry. 

Station 6019c. — ^Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. Lower part of lime-cemented soft gray to olive-colored 
limestone, with central parting of dark clay. The first hard, limy 
sandstone bed above the lower limestone and just above 60196. 
Culebra formation (upper part). Oligocene series. Collectors, 
D. F. MacDonald and T. W. Vaughan ; 1911. CaUianassa vaughani 
Kathbun (probably). CaUianassa ?, species. Hepatus^ species. 

Station 6019e. — ^Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. Third hard sandstone bed from bottom. Culebra forma- 
tion (upper part). Oligocene series. Collectors, D. F. MacDonald 
and T. W. Vaughan ; 1911. CaUianassa stridens Rathbun. 

Station 6012c. — ^Panama Canal Zone. Gaillard Cut. Top part of 
limy sandstone below upper conglomerate, near foot of stairs. 
Culebra formation (upper part). Oligocene series. Collectors, 
D. F. MacDonald and T. W. Vaughan; 1911. Natantia, family, 
genus, and species indeterminable. CaUianassaj species. CaUineotes^ 
species. Euryplax culehrensis Rathbun. 

Station 6020a. — Panama Canal Zone. Las Cascadas section, 
Gaillard Cut. Lowest fossiliferous bed. Third bed below lowest 
limestone beds separated by rows of nodules. Culebra formation 
(lower part of upper half). Oligocene series. Collectors, D. F. 
MacDonald and T. W. Vaughan; 1911. Balanus {Hesperiba- 
lanusf)^ sp. Axius reticulatvs Rathbun. CaUianassa avails Rath- 
bun. CaUianassa elongata Rathbun. CaUianassa crassimana Rath- 
bun. CaUianassa spinulosa Rathbun. CaUianassa quadrata Rath- 
bun. CaUianassa dbbreviata Rathbun. CaUianassa magna Rath- 
bun. GoniocJiele f armata Rathbun. Calappella quadrispina Rath- 
bun. CaUinectes reticulatus Rathbun. Thaumastoplax prima Rath- 
bun. Brachjrrhyncha, family, genus, and two species indeterminable. 

Station 6026. — ^Panama Canal Zone. About 200 yards south of 
southern end of switch at Bohio Ridge station, relocated line Panama 
Railroad. Foraminiferal marl and coarse sandstone. Culebra for- 
mation (upper part). Oligocene series. Collectors, D. F. Mac- 
Donald and T. W. Vaughan; 1911. CarpiUus^ species. 



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126 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Station 6019g. — ^Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. Fifth or topmost limestone. Emperador limestone. Oli- 
gocene series. Collectors, D. F. MacDonald and T. W. Vaughan; 
1911. Macrohrachiwm^ species. CaUianassa tenuis Rathbun. Mur- 
sia macdonaldi Eathbmi. PartTienope panamensis Eathbim. 

Station 6003. — ^Panama Canal Zone. Gatun section A, bed A (bot- 
tom of section). Gatun formation (lower part). Miocene series. 
Collector, D. F. MacDonald. Callianasm crassa Rathbun. 

Station 6029a, — Panama Canal Zone. One-fourth to one-half mile 
beyond Camp Cotton toward Monte Lirio. From lowest horizon in 
big cut. Gatun formation (lower part). Miocene series. Col- 
lectors, D. F. MacDonald and T. W. Vaughan; 1911. CaUianassa 
vaughani Rathbun. 

Station 60S3h. — Panama Canal 2iOne. Gatun section. Upper part 
of lowest bed. Gatun formation. Miocene series. Collectors, D. F. 
MacDonald and T. W. Vaughan; 1911. Lepas injvdicata Pilsbry. 
t Oatwnia proavita Rathbun. 

Station 6030. — ^Panama Canal Zone. One and one-half to 2 miles 
beyond Camp Cotton toward Monte Lirio. From 85-foot cut on 
north side of big swamp on relocated line, P. R. R. Gatun formation- 
Miocene series. Collectors, D. F. MacDonald and T. W. Vaughan : 
1911. Balanus concaws rariseptaiics Pilsbry. Callianassa vaughani 
Rathbun Gatunia proavita Rathbun. 

Station 6900. — Panama Canal Zone. Gatun Locks. Gatun forma- 
tion. Miocene series. Collector, D. F. MacDonald; May, 1911. 
Oatunia proavita Rathbun. 

Catalogue No. 113706^ Z7./S.iV.iI/.— Panama Canal 2lk)ne. Near Ga- 
tun. Gatun formation (?). Miocene series. Labeled "Miocene.'^ 
Collector, Rev. J. Rowell. Gatunia proavita Rathbun. 

Station 6659. — Panama Canal Zone. Near Gatun Dam. Gatun 
formation. Miocene series. Collector, one of the workmen; shipped 
by D. F. MacDonald ; 1911. Gatunia proavita Rathbun. 

Catalogue No. 136218^ U.S.N.M. — Panama Canal Zone. Gatun 
beds. Gatun formation. Miocene series. Collector, R. T. Hill* 
Callianassa hUU Rathbun. * 

Catalogue No. 135219^ f7./S.A^.il/.— Panama Canal Zone. Gatun 
beds. Gatun formation. Miocene series. Collector, R. T. HilL 
Mursilia ecristata Rathbun. 

Station 6S82Jc, — Costa Rica. Banana River; tenth fossiliferous 
zone below the uppermost one of the section. Probably equivalent 
to Gatun formation. Miocene series. Collector, D. F. MacDonald ; 
1911. EupKylax fortis Rathbun. 

Station 6882 j. — Costa Rica. Banana River; ninth fossiliferous 
zone below the uppermost one of the section. Probably equivalent to 
Gatun formation. Miocene series. Collector, D. F. MacDonald; 
1911. Euphylax callinectias Rathbun. 



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GEOLOGY AND PALEONTOLOGY OF THE QANAL ZONl^. 127 

StdHon 58821. — Costa Rica. Banana River; eighth fossiliferous 
zone below the uppermost one of the section. Probably equivalent 
to Gatun formation. Miocene series. Collector, D. F. MacDonald; 
1911. CaUin£ctes declivis Rathbun. 

Station 6882h. — Costa Rica. Banana River ; seventh fossiliferous 
zone below the uppermost one of the section. Probably equivalent to 
Cratun formation. Miocene series. Collector, D. F. MacDonald; 
1911. Leucoailia bananensis Rathbim. 

Station 6882g. — Costa Rica. Banana River; sixth fossiliferous 
zone below the uppermost one of the section. Probably equivalent to 
Gatun formation. Miocene series. Collector, D. F. MacDonald; 
1911. LeucosUia bananensis Rathbun. 

Station 6882 f. — Costa Rica. Banana River; fifth fossiliferous 
zone below the uppermost one of the section. Probably equivalent to 
Gatun formation. Miocene series. Collector, D. F. MacDonald; 
1911. LeucoaiUa bananensis Rathbun. 

Catalogs No. $£4287, U.S.N.M.—Coeta Rica. Moin Hill, near 
Liimon. Probably equivalent to Gatim formation. Miocene series. 
Collector, H. Pittier. Callianassa moinensis Rathbun. 

Station 6884d. — Costa Rica. Moin Hill; third fossiliferous zone 
below the uppermost; just abo^e level of the rails in railway cut. 
Probably equivalent to Gatun formation. Miocene series. Col- 
lector, D. F. MacDonald; 1911. Callianassa moinensia Rathbun. 

Station 6906a. — Panama Canal Zone. Chagres River, 50 to 75 feet 
below those of (17c) "5905" in lighter colored limestone according 
to incomplete evidence. Pliocene series. Collector, D. F. Mac- 
Donald ; May, 1911. Bcdanus glyptopoma Pilsbry. 

Station 6903. — Panama Canal Zone. From across Chagres River 
and probably 220 to 225 feet above level of river, top of hill opposite 
Alhajuela. Gray tufaceous limestone. Pliocene series. Collector, 
D. F. MacDonald; May, 1911. Balamis glyptopoma Pilsbry. 

Station 4269. — Costa Rica. City of Port Limon. Port Limon 
formation. Pliocene series. Collector, Dr. L. A. Wailes. Pa- 
chycheles latics Rathbun. Petrolisthea avitus Rathbun. OcHappa 
coataricana Rathbun. Heteractaea lunata (Milne Edwards and 
Lucas). Cardisoma gtuinhund Latreille. 

Station 6886. — ^Mexico. From the Sayula District of Chiapas. 
On the Arroyo Chapapoapam. Pliocene series. Collectors, Dr. 
C. W. Hayes and others, 1911. Balamis glyptopoma Pilsbry. 

Station 6038. — Panama Canal Zone. From black mud from lower 
end of Gatun Locks. Pleistocene series. Collector, D. F. MacDon- 
ald; 1911. Balanus ebumeus Gould. 



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128 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Station 6867. — Panama Canal Zone. From dark mud formation, 
about 10 feet above present sea level, near lower end of Gatun Locks. 
Pleistocene series. Collector, D. F. MacDonald; April, 1911. Ba- 
lanus ehumeus Gould. 

Station 5868, — ^Panama Canal Zone. From Mount Hope. Swamp 
ditch. Black mud formation. Pleistocene series. Collector, D. F. 
MacDonald; April, 1911. Balanua ehumeus Gould. 

Station 6850. — ^Panama Canal 2k)ne. Near Mount Hope in ditch 
through swampy ground. About one-fourth mile from present sea 
beach and about 6 to 8 feet above high tide. Pleistocene series. 
Collector, D. F. MacDonald; April, 1911. Macrohrachiwmt^ species. 
Nepkrops costatus Rathbun. Nephrops^ species. Axiusf, species. 
Hepatus ckilievms Milne Edwards. Calappa ftarrvmea (Herbst). 
LeucosUia jvrinei (Saussure). Leucosiidae, genus and species inde- 
terminable. Arenaevs^ species. Panopeus antepurpureus Kathbun. 
Panopeus tridentatus Rathbun. Eurytiv/m crenuUvtum Rathbun. 
Vca mxicTodactyluB (Milne Edwards and Lucas). Parthenope pleiR- 
t-ocenica Rathbun. 

In the following table the Cirripedia (see pp. 185-188) are in- 
cluded with the Decapoda. The letter " n " after a name in the first 
column indicates a new species or a new genus. The numerical head- 
ings refer to the same stations as in the above list but are arranged 
serially instead of chronologically. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 129 



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BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 131 

DESCRIPTIONS OF SPECIES. 

Subclass MALACOSTRACA. 

Order DECAPODA. 

Suborder NATANTIA- 

Family, genus, and species indeterminable. 

Plate 57, fig. 1. 

Locality, — ^Panama Canal Zone. Top part of limy sandstone below 
upper conglomerate, near foot of stairs, Gaillard Cut. Upper part 
of Culebra formation. Oligocene series. D. F. MacDonald and 
T. W. Vaughan, collectors. 1911. Station 6012t?. Cat. No. 324267, 
U.S.N.M. 

Material. — One specimen showing three segments from the pleon 
of a shrimp. Pleon compressed laterally. Each of the two overlap- 
ping segments has the posterior angle produced backward in a 
rounded lobe of moderate size. 

Family PALAEMONIDAE. 

MACROBRACHIUM, species. 
Plate 57, figs. 4 and 5. 

Locality. — Panama Canal Zone. Las Cascadas section, Gaillard 
Cut. From fifth or topmost limestone. Emperador limestone. Oli- 
gocene series. D. F. MacDonald and T. W. Vaughan, collectors, 
1911. Station 60195^. Cat. No. 324256, U.S.N.M. 

Material. — One propodus of left cheliped, minus finger. Slightly 
compressed, subcylindrical. Some of the outer crust is lacking, but 
in general, the segment widens rapidly for the proximal two-fifths, 
then widens gradually at the middle, but not at all in the distal two- 
fifths. There is no shallow sinus .in the lower margin behind the 
finger, as in M. jamaicense,^ M. acanthurus panamense ^ and others ; 
neither is the palm like that of M. mexicanum^'* which is not at all 
convex below, and has subparallel margins. 

The specimen resembles MaxyrobracKiwm more than it does any 
marine genus now existing in Panamian waters. 

^Coficer (AttootM) famaicensis Herbet, Natur. Krabben u. Krebee, vol. 2, 1792, p. 57, 
PU 27, flg. 2, 

*Ratlibnn, in SmltbsoD. Misc. Coll., toI. 59, No. 13, 1912, p. 1. 

*Palaemon mcaioanus Saussnre, Mtfm. Soc. Pbys. mst. Nat. Geneve, vol. 14, 1858. 
p. 468 [52], pi. 4, flgt. 27, 27a. 



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132 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Measurements. — ^Length of palm, 13 mm.; width, 4.3 mm.; thick- 
ness, 3.7 mm. 

MACROBRACHIUM?, ipmIm. 

Plate 57, fig. 9. 

Locality. — Panama Canal Zone. From near Mount Hope in ditch 
through swampy ground. About one-quarter mile from present sea 
beach, 6 to 8 feet above high tide. Pleistocene series. D. F. Mac- 
Donald, collector. April, 1911. Station 5850. Cat. No. 324248, 
U.S.N.M. 

Material. — One segment (perhaps the carpus) of the second or 
large pair of chelipeds, probably the left one. Subcylindrical, en- 
larging gradually to the distal end, slightly curved, a longitudinal 
row of 5 low conical spines irregularly spaced. 

Measurements. — ^Length, 9.5 mm. ; diameter, 1.7 mm. 

Suborder REPTANTIA. 

Tribe ASTACURA. 

Family HOMARIDAE. 

KEPHROPS COSTATUS. new ipMiM. 

Plate 57, figs. 13-17. 

Type-locality. — Panama Canal Zone. From near Mount Hope in 
ditch through swampy ground. About one-quarter mile from present 
sea beach, 6 to 8 feet above high tide. Pleistocene series. D. F. 
MacDonald, collector. April, 1911. Station 5850. 

Types.— Cat No. 324246, U.S.N.M. 

Material. — ^Three dactyli of left cheliped, one of which is fairly 
complete and is taken as the holotype; the other specimens show 
only the distal half or two-thirds. A fourth specimen (distal half 
only) represents a fixed finger perhaps and if so belongs on the left 
side. 

Holotype. — Length 9.5 mm. In dorsal view the inner or right mar- 
gin is sinuous, the tip curved strongly inward; viewed from the 
inside, both edges are sinuous, curving downward toward the tip. 
Upper and lower surfaces a little flattened. Five longitudinal 
costae, 2 dorsal, 2 ventral, 1 inner; each costa marked by a line of 
fine granules, with a row of punctae adjacent. On the proximal half 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 133 

there is some intercostal granulation. Prehensile edge armed with 
fine teeth and divided into 3 sinuses separated by 2 large teeth ; the 
distal of these has its distal edge normal and its proximal edge 
oblique to the margin of the dactylus ; the top of the other large tooth 
is broken off; the terminal bay has a somewhat enlarged, but still 
small, tooth at its middle. 

Paratypes. — {a) Distal half of dactylus, but with small tip lack- 
ing, same width as holotype; terminal sinus same length but more 
curved, so that the distal border of the boundary tooth is shorter; 
middle sinus half as long, nondentate, next boundary tooth broader 
than in holotype. 

(6) Dactylus with proximal end lacking, same width as holotype, 
costae more rounded, terminal sinus a little shorter, boundary tooth 
with end missing, enlarged middle tooth better developed than in 
holotype, pointing obliquely distad; middle sinus longer, boundary 
tooth broken. 

(c) Propodal (?) finger broader than the others, showing one 
sinus nearly equal to 2 sinuses of the holotype and limited by a large 
tooth with nearly equal sides. 

I have placed this species in Aephrops on account of the ribbed 
fingers irregularly toothed. The variations in the dactyl may repre- 
sent either individual or sexual variation. 

NEPHROPS, ipccies. 

Plate 57, fip^. 25 and 26. 

Locality. — Panama Canal Zone. From near Mount Hope in ditch 
through swampy ground. About one-quarter mile from present sea 
beach, 6 to 8 feet above high tide. Pleistocene series. D. F. Mac- 
Donald, collector. April, 1911. Station 5850. Cat. No. 324249, 
U.S.N.M. 

Material. — Dactylus of right cheliped, 12 mm. long; distal half 
moderately curved toward the propodal finger, but the whole finger 
strongly curved downward; 6 strong, longitudinal costae, 3 dorsal^ 
1 marginal, 2 ventral ; about 9 lines of punctae ; the prehensile teeth, 
36 in all, are larger and more projecting in that two-fifths of the 
margin just posterior to the middle. 

After the above description was written the proximal half of the 
specimen was accidentally crushed and destroyed. 

Although the dactylus is much more curved than in any species of 
NephropSj yet its ornamentation is so similar to that of the preced- 
ing species, N. costcUus^ that it is referred to the same genus. 



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134 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Tribe ANOMURA. 

Superfamily GALATHEIDEA- 

Family PORCELLANTOAE. 

PACHTCHELES LATU8, new spmIm. 
Plate 57. flj^. 21-23. 

Type-locality. — Costa Bica; Poii: Limon. Pliocene series. Dr. 
L. A. Wailes, collector. Station 4269. Holotype, left manus with 
propodal finger; inner proximal corner of manus broken oflF. Para- 
type, left manus, with both fingers; proximal portion of manus 
broken oflF. 

Type.— Cat. No. 324264, U.S.N.M. 

Measurements. — Width of palm, 4.6 mm. ; length of same to sinus, 
5.1 mm. ; length to end of finger, 6.7 mm. ; greatest thickness, 2.3 mm. 

Holotype. — Outer and inner margins thick and strongly curved 
in dorsal view; upper surface coAered with granules crowded to- 
gether and of varying size ; the granules are continued on the outer 
surface and a little way on the imder surface ; they are then replaced 
by squamiform granules and short rugae which are continued over 
the inner surface. There are no marginal lines indicated. At the 
distal end, the width from the articulating condyle to the inner angle 
is nearly as great as to the outer margin. The fixed finger is short 
and stout, width subequal to length; a bit of the tip is, however, 
missing; a low tooth occupies the greater part of the basal half of 
the prehensile edge. 

Pa/ratype. — Smaller than the holotype and much worn so that the 
granulation is not well marked. Tooth at base of immovable finger 
minute. Movable finger very short and broad, granulate, with a 
basal prehensile tooth, its surface granulate. 

In general shape and granulation, this form resembles the manus 
of the Recent P. grossimanus (Gu^rin) from Peru and Chile, but in 
the latter the outer margin is paved with larger granules forming a 
definite edge, and the propodal finger is longer and more curved. 

PETROLISTHES AVITUS. new ipedM. 

Plate r>7, fi-s. 18-20. 

Type-locality. — Costa Rica; Port Limon. Pliocene series. Dr. 
L. A. Wailes, collector. Station 4269. 

Type.—Cni. No. 324266, XT.S.N.M. 

Holotype, — Palm of left cheliped, showing the greater part of the 
upper and lower surfaces including the inner margin and the distal 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 135 

articulating edge of the- lower surface. Outer edge, proximal end, 
and finger missing. Upper surface covered with coarse granulated 
striae of very different lengths, varying from 1 to 10 or 12 granules, 
and arranged obliquely longitudinally. Lower surface covered with 
curved, wavy and punctate striae starting almost at right angles with 
the inner margin, curving slightly forward and then abruptly back- 
ward; so that the greater part is more longitudinal than transverse; 
the striae are somewhat subdivided and followed outwardly by 
shorter striae; at the inner end they terminate abruptly, so that from 
above they have the appeal ranee of 13 trimcated shallow teeth. 
Length 5.2 mm. 

This manus resembles that of two common recent species, P. 
amuUu8 (Gibbes)', and P. gaZatkinus (Bosc)*, both found on the At- 
lantic as well as on the Pacific side of the continent. The upper 
surface of the palm is similar in P, armatvs^ that is, it is ornamented 
with short, irregular striae, which are, however, parallel to the inner 
margin, while in the fossil form they diverge proximally from the 
margin. The lower surface of P, avitus^ on the otlier hand, resembles 
more closely that of P. gdlathinus^ but in the latter, the striae trend 
more strongly forward on leaving the inner margin, and that margin 
itself is not formed of such strongly marked teeth. 

Superfamily THALASSINIDEA. 
Family AXIIDAE. 

AXIUS BBTICULATUS. new spmIm. 
Plate 57, figs. 2 and 3. 

Type-locality. — ^Panama Canal Zone.' Las Cascadas section, Gail- 
lard Cut. From lowest fossiliferous bed. Third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a. 

Holotype.—Cat No. 324260, U.S.N.M. Left propodus of first 
pereiopod, embedded in a nodule and showing the finger and the 
greater part of the palm, except the proximal end and the distal 
upper corner. An impression of the same is shown in another piece 
of the nodule. The segment as uncovered is 14.3 mm. long, greatest 
height 5 mm., length of finger 7 mm. The palm is greatly swollen 
and at the top rounds over into a broad upper surface about 2.4 mm. 
in width. The shell is considerably cracked and in life may not have 
been as thick as it appears. The lower margin is sinuous, forming a 
bay at about the distal third of the palm ; so far as the edge is pre- 
served it is formed of small bead granules. The outer and upper 
surface is ornamented with granules irregular in size and disposi- 

» Porcellana armata Gibbes, Proc. Amer. Assoc Adv. Scl., vol. 3, 1850, p. 100. 
• Porcellana galathina Rose, Hist. Nat. Crust., vol. 1, 1802, p. 233, pi. 6, Hg. 2. 



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136 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

tion, larger and thicker on the distal part midway between upper 
and lower margins, elsewhere smaller and to a large extent forming a 
reticulate pattern ; proximally on the upper part of the outer surface 
the raised reticulate lines are smooth, or non-granulate. 

The outer surface of the finger is rather regularly tapering, the 
lower margin directed slightly upward, the superior margin nearly 
straight; surface smooth; finger thick, the upper surface oblique or 
beveled; traces of fine teeth are visible on the prehensile edge. 

AXIUS?, ipeaes. 

Plate 57, fig. 10. 

Locality. — Panama Canal Zone. From near Mount Hope, in ditch, 
through swampy ground. About one-quarter mile from present sea 
beach, 6 to 8 feet above high tide. Pleistocene series. D. F. Mac- 
Donald, collector. April, 1911. Station 5850. Cat No. 324250^ 
U.S.N.M. 

Material. — ^A single movable spine, 6.7 mm. long, with the tip 
broken off, resembles the styloid scaphocerite or movable acicle of the 
outer antenna of some species of Axius. The spine is somewhat 
8-angled, the most acute edge being dorsal, the two blunt edges being^ 
nearer together and ventral. There are a few punctae : 4 large ones 
in a row on tiie ventral surface; 2 large, external, far apart, just 
below the upper margin; 4 small ones, internal, 3 of which form a 
triangle near the middle, while the other is nearer the distal end. 

Family CALLIANASSIDAE. 

KEY TO THE SPECIES OF CALLIANASSA HEBE DESCRIBED. 

The material is insufficient to distinguish between the larger and 
the smaller chela of the same pair, which also may vary in shape and 
size in the two sexes. 
A\ Manus and carpus meeting in an oblique line. 

B*. Lower margin of manus serrated oihjZw, p. 13T 

B*. Lower margin of manus smooth lacwMsa, p. 13S 

A". Manus and carpus meeting in a vertical line. 
B\ Lower margin of manus directed forward and upward, at least in part. 
G\ Palmar portion of manus distinctly longer than high. 
D\ Palm compressed. 

E*. Palm elongate; margins strongly convergent eion(/ata, p. 13^ 

E*. Palm less elongate ; margins moderately convergent — scottiy p. 140 

D*. Palm swollen moinensiSy p. 142 

C*. Palmar portion of manus about as long as high, or shorter. Upper 

margin of manus directed forward and downward toward the lower 

margin. 

D\ Immovable finger very thin, a cross-section near Its base being more 

than twice as long as wide. A strong tooth In the sinus between the 

fingers or on the base of the Immovable finger apinulosa, p. 145 

D*. Immovable finger thicker, a cross section near Its base being less 
than twice as long as wide. A tooth In the sinus between the fingers 
but near the base of the dactylus tenuis, p. 144 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 137 

A". Manns and carpus meeting In a vertical line — Continued. 

B*. Ix)wer margin of menus directed straight forward or nearly 8o; lliat is, 
at right angles to its proximal margin. 
C. Upper margin of manus subparallel to lower margin. No tooth in 
sinus between fingers. Carpus much higher than long. 

D*. Lower margin serrulate gwodraf a, p. 145 

D*. Lower margin granulate ^toi^Zai, p. 146 

C*. Upper margin of manus directed forward and downward toward lower 
margin. 
D\ A large tooth in sinus between fingers and situated on base of im- 
movable finger abbreviatay p. 147 

D*. A small tooth in sinus between fingers, and situated near movable 

finger. Carpus very little, if at all, higher than long AiUt, p. 148 

D*. No tooth in sinus between fingers. Fingers long and strong. 

vauffliani, p. 148 
B*. Ijower margin of manus unknown. A stridulatiug ridge near the hori- 
zontal upper margin atridens, p. 161 

A'. Meeting of manus and carpus unknown. 

B\ Immovable finger slender. Distal articulating edge of manus crenulate 

and very oblique cra^^imana, p. 141 

B^ Only the dactylus known. 

C*. Dactylus of large size. Cross section at base subcylindrical. Pre- 
hensile edge thin 1 niagnay p. 151 

C*. Dactylus half as long as preceding, more compressed. Prehensile 
edge thicker craasUf p. 152 

The 2 Callianassas to which specific names are not given are ex- 
cluded from the above key^ 

CALLIANASSA OVALIS^ new ipftdei. 
Plate 50, figs. 1-4. 

Type-locaUty. — Panama Canal 2k)ne. Las Cascadas section, Gail- 
lard Cut From lowest fossiliferous bed, third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a. Holotype 
and one paratype. Cat. No. 324269, U.S.N.M. 

Holotype. — ^A left chela with wrist and arm attached, and enclosed 
in a nodule which is split in two. The specimen had been crushed, 
and the half nodule containing the impression shows also small 
pieces of the cheliped itself. Wrist and manus together oblong- 
oval ; line between them oblique ; upper and lower margins of manus 
slightly convex; greatest width about equal to the upper length; 
from the widest point, the lower margin of the propodus slants 
upward. Fingers directed straight forward and of subequal length. 
Immovable finger an isosceles triangle, the base of which is two- 
^rds as long as either side; end blimt; cutting edge with a shallow 
triangular tooth at the proximal two-fifths. Movable finger sub- 
oblong, end broad, cutting edge with a broad, shallow, rounded tooth 



Digitized by VjOOQIC 



138 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

near the base. The merus appears to be about twice as long as high 
and is dilated at the middle. 

Measurements. — Length of carpus and propodus, measured from 
lowest point of articulation with merus to end of propodal finger, 
20.5 mm.; proximal width of propodus (approx.), 9.4 mm.; greatest 
width of same, 10 mm.; distal width of same, 8.7 mm.; superior 
length of same, 10.2 mm. ; inferior length of same, 13.2 mm. ; length 
of propodal finger measured on cutting edge, 5.7 mm.; length of 
dactylus, 5.7 mm. 

Paratype. — (a) Left manus with part of the fixed finger, em- 
bedded in a nodule and exposing the outer surface. Larger than 
holotype. The lower margin of the manus has blunt serrations 
resembling the stumps of spines; above the margin is a row of five 
distant punctae; at the distal end there are two rows of granules 
leading toward the upper part of the fixed finger. 

Paratype, — (6) Carpus of left cheliped embedded in a nodule and 
imprint of same, showing the distal and upper margins and a large 
])art of the outer surface. The distal margin is oblique and concave 
and has a little rounded lobe at each end ; the upper margin is arcuate 
and has a submarginal groove. 

CALUANASSA LACUNOSA, new apeciM. 
Plate 59, tips. 6-11. 

Type-locality. — One-quarter mile south of Empire Bridge, Canal 
Zone, Panama; from lower dark clay beneath lower conglomerate. 
Lower part of Culebra formation. Oligocene series. D. F. Mac- * 
Donald and T. W. Vaughan, collectors. 1911. Station 6012a. Two 
specimens, each a left propodus of the first cheliped. Cat. No. 
324278, U.S.N.M. 

Holotype. — ^The palmar portion of a propodus, the fixed finger 
broken off near its base; the propodus is incomplete near the dactylus 
and also at the proximal end ; this end is, however, fairly complete 
on the inner surface, so that the measurements may be stated with 
approximate accuracy. Length, at the level of the sinus between 
the fingers, 16 mm. ; greatest height, 15.6 mm. ; least height, 15.2 mm. ; 
thickness, 6.6 mm. The upper margin is a little arched, the lower 
nearly straight ; they converge a little distally ; they are very thin ; 
the top of this thin edge along the upper margin is set with oblong 
tubercles, dorsal in position; the lower edge is smooth. The outer 
surface has a row of 6 pits a little above the edge and fairly evenly 
spaced ; they have a raised edge and are distally inclined, indicating 
that they were sockets for hairs or bristles; 3 similar pits far apart 
are close up under the upper margin; of the few scattered pits, 3 
form a triangle at the distal end, the one near the base of the fixetl 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 139 

finger being the largest; there is also a medium-sized pit a little 
helow the middle, while 3 small ones are visible near the proximal 
end. A prominent blimt ridge runs from near the top of the fixed 
finger obliquely backward and upward for a short distance on the 
palm; near this ridge, but chiefly above it, and partly behind it, 
there is a patch of low tubercles, mostly oblong but very irregular 
in shape and size. On the inner surface there is a row of 8 sockets 
a little distance below the thin, upper, marginal rim ; these sockets, 
while of good size, have very narrow, perpendicular openings. Above 
the lower margin there is a row of 12 sockets; this row distally ap- 
proaches close to the edge, while the sockets themselves become larger 
and farther apart; they are very oblique to the margin and are 
ahnost more distal than lateral in their inclination. These 2 rows 
of sockets on the inner surface are more prominent than any others; 
below the middle there are 10 or 12 scattered sockets mostly small; 
near the middle there is a patch of tubercles, somewhat masked by 
a thin layer of adhering matrix. On the inner surface there is a 
blimt ridge leading back from the finger similarly placed to that on 
die outer surface, but lower and wider. The finger is slender, judging 
from the section at its base. 

Paratype. — ^This propodus shows the outline of the proximal end, 
but the distal end is broken off and not a vestige of the finger remains. 
The size is less than in the holotype : length, at the level of the sinus 
between the fingers, 10.3 mm.; greatest height, 11.4 mm.; least 
height, 10 nun.; thickness, 5.2 mm. The margins are more con- 
vergent than in the holotype and the palm is relatively shorter. The 
upper as well as the lower edge is almost smooth. On the outer sur- 
face the sockets in the upper submarginal row are 4, as there is an 
additional one visible at the proximal end; the sockets of the lower 
row are fewer and more distant than in the larger specimen, as only 
5 can be detected ; of the scattered sockets, 3 form a distal triangle, 
while 10 or 12 small ones are disposed transversely near the proximal 
end; the protuberances above the oblique ridge leading from the fixed 
finger consist of a few small granules. On the inner surface, a 
row of 8 submarginal sockets above, as in tlie holotype; near the 
lower margin only 10 sockets can be counted, because the distal 
comer is broken away ; scattered sockets 16 or 18, below the middle ; 
a few granules close to the sinus between the digits. 

CALLIANASSA ELONGATA. ii«w ipedM. 
Plate 60, figs. 4-6. 

Type'localify. — ^Panama Canal Zone. Las Cascadas section, 
Gaillard Cut. From lowest fossiliferous bed, third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
8370*— IS— BulL 103 ^10 



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140 bullbun rL08, unitbd states national museum. 

half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a. 

Holotype.—C2it No. 324271, U.S.N.M. A left manus, with base 
of immovable finger attached; this shows all of the outer surfacey 
except the margin bordering the dactylus ; nearly the whole of the 
inner surface is concealed by the matrix. The upper margin is nearly 
straight throughout its length, and is bluntly angled ; lower margin 
very sinuous, the manus being very much deeper in its proximal 
than its distal half, edge serrulate or spinulous throughout its length ; 
proximal edge vertical, nearly as long as upper margin. On the 
outer surface there is a group of granules at the distal two-sevenths 
and just below the middle; there is a curved row of granules near 
the sinus, which is continued upon the propodal finger near its upper 
edge ; 6 granules in all are visible. Finger very slender, inclined 
downward. The dactylus must have been very stout, and the adja- 
cent edge of the manus very oblique, but it is now incomplete. 

Measixrements. — Superior length of manus (approx.), 15.7 mm.: 
length to sinus, 20.4 mm. ; greatest height, 16.2 mm. ; proximal height 
(approx.), 15.1 mm.; distal height, 12.4 mm. 

CALUANASSA SCOTTI Brown mud PUsbry. 

Plate 00, figs. 9-12. 

Callianassa scotti Brown and Pilsbry (part), Proc. Aead. Nat. Sol. Phila., 
vol. 64, 1913, p. 503, pi. 22, figs. 1 and 3 (not fig. 2). 

Locality. — Panama Canal Zone. From the 4 feet of dark strati- 
fied tuff and clay immediately overlying the lower limestone bed. 
Las Cascadas section. Upper part of Culebra formation. Oligo- 
cene series. D. F. MacDonald and T. W. Vaughan, collectors. 1911. 
Station 60196. Cat. No. 324279, U.S.N.M. 

MateTial, — A left manus, removed from the matrix, and incom- 
plete at the proximal end. Outer surface very convex from upper to 
lower margin ; upper margin straight, with a blunt marginal line ; 
lower edge acute and serrulate with fine appressed teeth or spines^ 
inclined upward toward the base of the finger, then downward : 
three granules or tubercles in a curved row just outside the edge of 
the sinus between the fingers. The cross section of the finger near 
its base is very small, in relation to the manus. Inner surface 
slightly convex, from upper to lower edge, and with a depression on 
either side of the base of the fixed finger; numerous granules near 
the distal end, some arranged in a curved band between the two 
fingers. 

Measurements, — Greatest height of manus, 25.2 mm. ; least height 
of same, 23.3 mm. ; length of same to digital sinus, 24.6 mm. ; thickness 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 141 

of same, 10.9 mm. ; greatest diameter of fixed finger near its base, 
6.5 mm. ; least diameter of same, 4.2 mm. 

Holotype and paratypes. — ^In the material included by Mr. Brown 
and Dr. Pilsbry under C. scotti are specimens of three species; two 
of these species were figured with the original description; the larger 
species illustrated ^ by figures 1 and 3, plate 22, is chosen to bear the 
name scotti^ and the original of figure 1 may be designated as the 
type-specimen, as it is free from the matrix and shows more of the 
characteristic granulation than does the original of figure 3, which 
is furnished with a propodal finger, but is half embedded in a matrix. 
It also shows, although roughly, a row of 5 pits on the upper margin ; 
this row, however, slopes downward a little distally on to the inner 
surface. The greatest height of the type is 27 mm., length at level of 
, sinus between fingers, 28 mm. 

Still a third specimen, also a left manus, was taken at the same 
place ; the surface is much worn, but the proximal angles are well de- 
fined. The typo-locality is in the lignitic layers, about 65 feet below 
the base of the Pecten bed at Tower N., Las Cascadas section, Gail- 
lard Cut; Prof. W. B. Scott, collector, 1911. Type, Cat. No. 2259, 
Mus. Acad. Nat. Sci., Philadelphia. 

The hand mentioned by Brown and Pilsbry ,2 as collected by W. M. 
Gabb in Costa Bica, apparently belongs to C. scotti. It shows well 
the upper marginal row of pits, numbering 7, which drops disbally 
on to the inner surface. The specimen is labeled " Miocene." Cat. 
No. 2255, Mus. Acad. Nat. Sci., Philadelphia. 

. CALLIANASSA CRASSmANA* new tiwdet. 
Plate 61, figs. 15-17. 

Type-locality. — Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. From lowest fossiliferous bed, third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a. 

Measurementa. — Distal height of palm, 22 mm. ; thickness of palm, 
at least 7.7 nmi. ; length of crenulated lobe bordering dactylus, 10.2 
mm.; upper length of dactylus (tip broken off), 18 mm.; greater 
diameter at the break near the tip, 1.7 mm.; lesser diameter at same 
point, 1.3 nmi. ; greater diameter at a break about middle of finger, 
8.8 mm. ; lesser diameter at same point, 2.8 mm. 

Holotype.— CB^t No. 824273, U.S.N.M. A portion of the distal end 
of the left manus with the propodal finger attached ; embedded in a 
nodule. This must have been a very large specimen. The oblique 

> Ptoc Acad. Nat. 8lc.» PhiUu, voL 64, 1918, p. 508 * Idem, p. 604. 



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142 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

margin of the lobe of the manus which overlapped the dactylus is 
very oblique and is straight and deeply crenulated with about 16 
crenules; the length of the lobe is as great as the distance across the 
adjoining sinus and the propodal finger. This sinus is very deep 
and proximally subtruncate. The immovable finger is very narrow in 
relation to the size of the palm; it is slightly compressed, anfi bends 
upward; on the inner surface just within the lower margin there is 
a row of pimctae ; on the outer face there are 3 larger spots, 2 near 
the lower margin and one near the sinus, which may be sockets for 
setae. A tubercle on the outer surface just behind the sinus. The 
inner surface of the manus is deeply channeled out near the sinus 
between the fingers. 

The holotyi)e is in two pieces, as the immovable finger is broken in 
two, the distal portion embedded in that half of the nodule which 
bears the imprint of the remainder of the holotype. 

This species is very near C. scotti^ but has a larger sinus between 
the fingers, a longer propodal finger, and a shallower sinus in the 
lower margin of the manus just behind the finger. 

CALUANASSA MOINENSIS» new species. 

Plate 60, flgs. 1-3. 

Type-locdlity. — Moin Hill, near Limon, Costa Bica. Probably 
equivalent to Gatun formation. Miocene series. H. Pittier, collector. 
Propodus of right cheliped, and an impression of half a finger. 

Measurements. — Length (approx.) of manus, to sinus between 
fingers, 9 mm. ; height of same, 7 mm. ; thickness of same, 4.7 mm. ; 
length of immovable finger (tip broken off), 8.6 mm. 

Holotype. — Cat. No. 324287, U.S.N.M. Palm much swollen, cross 
section ovate, lower margin viewed from the side very arcuate, upper 
margin slightly so. The surface has almost entirely lost the outer 
white layer, but the next layer is gray and is crossed transvei-sely by 
many very short rugae, which are strongest on the lowest part of the 
outer surface. There is a distinct line below dividing the inner from 
the outer surface and marked by an irregular row of very fine 
punctae. On the inner surface considerably below the upper margin 
there is a row of large punctae. 

The immovable finger is slender, bent downward and curved 
inward. It has 7 more or less defined ridges, the bluntest of which 
is the most inferior; either side of the ridge representing the pre- 
hensile edge there is a granular ridge, the outer of which is less 
elevated ; in addition, there are 2 ridges on the outer surface and one 
on the inner; near each ridge there is a row of fine punctae. Pre- 
hensile edge armed with small irregular teeth. There is an unusually 
deep furrow above tiie principal ridge on the inner surface. 



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GEOLOGY AND PAI-EONTOLOGY OF THE CANAL ZONE. 143 

Paratype (a).— Cat. No. 324287, U.S.N.M. On a small piece of 
crumbling rock there is an impression of the distal half of a finger, 
but not the finger of the holotype. The impression has a similar 
curvature and 3 rows of punctae are present. 

Paratype (&).— Cat. No. 324288, U.S.N.M. Moin Hill, Costa Rica; 
third fossiliferous zone below the uppermost; just below level of rails 
in railway cut. Probably equivalent to Gatun formation. Oligocene 
period. D. F. MacDonald, 1911. Station 5884<i. A right propodus 
without finger, similar in size to the holotype and corresponding in 
its characters. 

CALUANASSA 8PINULOSA, new ipedeik 

Plate 61, figs. 6-9. 

Cailianaasa scotti Pilsbry (part), Proc. Acad. Nat. Sci. Phlla., vol. 64, 1913, 
p. 503 (not pi. 22, figs. 1-3). 

Type-locality. — ^Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. From lowest fossiliferous bed, third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald and 
T. W. Vaughan, collectors. 1911. Station 6020a. 

Holotype.— C2X. No. 324272, U.S.N.M. Left propodus which was 
encased in a nodule; the manus has been removed except the proximal 
part of the outer surface; of the propodal finger only the impression 
remains. Length of manus less than greatest height ; the upper and 
lower margins converge distally, the upper margin convex, the lower 
sinuous. Outer surface very convex in a vertical direction, having 
a few scattered granules, also a short vertical line of 3 granules near 
the upper distal corner, a row of 6 distant granules just above the 
lower margin, and a sharp granule near the sinus. The lower margin 
is very thin and serrated; between the serrations are the truncated 
bases of movable spines; 9 such spines remain. Upper edge bluntly 
margined. Inner surface convex except near the inferior and distal 
margins ; a row of 3 granules on the middle line, 2 above the base of 
the propodus, 2 parallel to the lower margin, and many small granules 
just above that margin. An obliquely longitudinal line of pits below 
the upper margin. 

The propodal finger is very much smaller than the dactylus and 
thin; a cross section near its base is somewhat diamond-shaped, the 
impression of the thumb (viewed sideways) is subtriangular, end 
curved upward; prehensile edge concave; at its base in the sinus be- 
tween the fingers there is a short but strong curved tooth; on both 
outer and inner surfaces of the thumb, leading down from the palm, 
there is an oblique ridge. 

Measurements. — ^Length of propodus to end of finger (approx.), 
22.4 mm. ; fishgth of manus, measured on the inner side, to sinus, 14.7 



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144 BULLETIN 103, UNITED STATES NATIONAL MUSEUM, 

mm.; greatest height (proximally), 15.3 mm.; least height (distally), 
13.3 mm. 

Pardtypes. — {a) One left propodus with about half of the fixed 
finger attached was taken in the lignitic layers, about 65 feet below 
the base of the Pecten bed at Tower N., Las Cascadas section, Gail- 
lard Cut; central part of Culebra formation, Oligocene series; Prof. 
W. B. Scott, collector; 1911 (Mus. Acad. Nat. Sci. Philadelphia). 
The specimen is half embedded in a nodule which conceals the inner 
surface and the upper part of the outer surface. It is nearly as large 
as the type. It shows the large tooth at the proximal end of the pre- 
hensile edge of the fixed finger, the tubercle on the outer surface near 
the sinus between the fingers, and roughly, the ornamentation on the 
lower edge of the palm. 

(6) From the same source, a similar left propodus half embedded 
in a nodule but so as to expose the inner surface and the upper edge ; 
it shows the characteristic row of pits on the inner surface just below 
the upper edge, the row sloping downward distally. 

(c) Also a third specimen, free from the matrix but with the edges 
broken ; it shows the palmar ridge leading to the fixed finger and the 
basal tooth on the edge of the latter. 

CALUANASSA TENUIS, new ipedcs. 

Plate 60, figs. 13 and 14. 

Type-locality. — Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. From fifth or topmost limestone. Emperador limestone. 
Oligocene series. D. F. MacDonald and T. W. Vaughan, collectors, 
1911. Station 6019pr. 

H olotype.— C^t. No. 324282, U.S.N.M. Left manus with propodal 
linger broken off, segment shorter than its greatest height, in general 
subrectangular, with upper and lower margins converging distally; 
outer surface very convex from top to bottom; upper and lower 
fdges marginate, the lower one very thin; inner surface convex ex- 
cept near the bottom where it is flat, at the proximal tod where there 
is a furrow parallel with the articulation, and near the distal sinus 
where there is a depression. The propodal finger is very much 
smaller than the dactylus, and its cross section is suboval with 
pointed ends ; the sinus is about as wide as the thumb ; it bears, close 
up to the insertion of the dactylus and on the edge of the outer sur- 
face, pointing forward, a short, broadly triangular spine. Surf acas 
much worn, so that it is difficult to tell whether unevennesses are 
natural or not. 

MecLSurements. — Length of manus, measured to sinus, 11.4 nun.; 
greatest height, 13 mm. ; distal height, 10.3 mm. ; thickness 5.3 mm. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 145 

CALUANASSA QUADRATA, n«w species. 

Plate 62, figs. 4-14. 

CalUanassa scotti Brown and Pilsbby (part), Proc. Acad. Nat. Sci. Phila., 
vol. 64, 1913, p. 503, pi. 22, fig. 2 (not figs. 1 and 3). 

Bepresented by 2 specimens of the left manus from which the 
propodal finger has been broken off. The smaller one is used as 
the type, as it is the better preserved. 

Type-locality. — Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut From lowest fossiliferous bed, third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a. Holotype 
And paratype. Cat. No. 324275, U.S.N.M. 

Measurementft (In mm.). — 

Holotype. Paratype. 

Height of manus near middle 13.3 15.3 

Length of manus measurerl to sinus 12. 8 15. 7 

Thickness of manus 6. 2 7.'5 

Holotype. — Hand squarish, with the corners rounded off. The 
upper and lower margins are subparallel, the upper slightly arched, 
the lower with a shallow distal sinus. Distal margin, at the articula- 
tion with the dactylus, nearly vertical. Outer surface slightly con- 
vex in a longitudinal direction, strongly convex vertically, furrowed 
at the distal end across the middle third; inner surface moderately 
convex except at the distal and lower portions, where it is concave; 
there is a groove just above the lower margin which widens as it ap- 
proaches the finger. Lower margin serrulate ; upper margin bluntly 
angled except in the distal third, where it is rounded; just within 
this margin there is a row of sockets of which 3 can be made out. 
On the outer surface near the sinus between the fingers there is a 
tubercle and near the carpus 3 granules far apart in a vertical row ; 
on the inner surface there is a row of granules, running almost longi- 
tudinally near the middle and thence downwards toward the sulcus 
between the fingers; 2 tubercles near the articulation with the dac- 
tylus. Propodal finger narrow, much compressed, subtriangular at 
tie base in cross section, with the small end of the triangle down. 
Base of dactylus very large. 

Paratype (a). — ^Larger than the type; upper margin straighter; 
the tubercle on the outer surface near the sinus is of good size ; on 
the inner surface near the middle there are numerous granules instead 
of the single row in the holotype ; 2 tubercles near the dactylus. 

Doubtful specimen. — ^A single specimen of a left carpus, from the 
same locality as the types may belong to the same species. The 
inner surface is mostly concealed by the matrix. Outer surface 



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146 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

very convex from the upper to the lower margin ; convexity akin to 
that of the manus of this species, which is the reason for placing 
the carpus here rather than with OaUianassa elongata or any other 
species occurring at the same locality. Carpus about If times as 
high as its greatest width, which is in the upper part; the angle 
formed by the superior and the distal margins is a little less than a 
right angle ; superior margin straight ; distal angle projecting above 
the articulation with the manus. The inferior distal angle projects 
even more below the articulation; the angle is obtuse; from it the 
margin rounds downward and then upward in a single curve; the 
lowermost part is finely serrate. 

Measv/remenU. — ^Height of carpus, between distal angles, 18.S 
mm. ; greatest height, 18.8 mm. ; greatest width, 12.8 mm. ; width on 
upper margin, 12 mm. ; greatest width below the articulation with the 
merus, 11.8 mm. 

Four specimens before me from the collection of the Academy of 
Natural Sciences of Philadelphia belong to this species; they are one 
right and three left chelae and form part of the material referred by 
Brown and Pilsbry to their C, scotti. (Paratype h) One left chela 
is that figured on plate 22, fig. 2.* All are larger than the type ma- 
terial described above, but so far as their characters are preserved 
they agree in essentials with the type. (Paratype c) The right chela 
(the largest specimen) its about 23 mm. long by 21.2 mm. wide, and 
possesses a longer piece of the propodal finger than the other speci- 
jmens ; the exposed cross section of the finger is oval. (Paratype d) 
The shortest of the left chelae has a large part of the outer layer pre- 
served on the infero-distal and inferior surface, where it is covered 
with granules, arranged without regularity except for a row on the 
outer surface parallel and close to the lower edge. This row is not 
visible in my figure 14, plate 62. The longest of the left chelae 
(Paratype e) is about 24 mm. by 19 mm., that is, considerably nar- 
rower than (c). 

CALLIANASSA TOULAI. new ipeciei. 

" Krnbbenscheren " Toula, Jahrb. der k. k. Geolog. Heichsanstalt, vol. 61, 
1911, p. 512 [26], pi. 30 [11, fig. 14. 

Founded on two chelae, one with palm 20.6 mm. long, 16.5 mm. 
wide, the other (without movable finger) with palm 9.4 mm. long, 
5.5 mm. wide. 

Outer surface of palm arched, inner surface almost flat and with a 
flat depression close to the lower margin and extending from the 
fixed finger. The upper and lower margins are sharp, the former 
bears 3 spinelike projections directed forward, the lowest spine ends 

^ Proc. Acad. Nat. Scl. Phlla., vol. 64, 1918, p. 508 



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QBOLOGY AND PALBONTOLOOY OF THE OANAL ZONE. 147 

in a sharp margin ; lower margin very finely and sharply granulate 
to the finger tip. Outer surface smooth and glossy, thickly covered 
near the lower margin with fine granules. Inner surface covered 
with very small, oblong punctae. A stout tooth on the prehensile 
margin of the immovable finger visible from the outside; another 
^oth on the movable finger, visible from the inside. 
Type-locality. — Gatun. Miocene series. 

CALLIANASSA ABBREVIATA, new species. 

Plate 63, figs. 1-6. 

Type-locality. — Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut From lowest fossiliferous bed; third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020<^. 4 specimens. 

rype^.— Cat. No. 324274, U.S.N.M. 

Measurements. — ^Length of manus to sinus, 10.3 mm.; greatest 
height, 10 mm.; distal height, 8.4 mm. 

Holotype. — Spechnen of right manus partially embedded, showing 
inner surface and half of outer surface; an imprint of the inner sur- 
face of the holotype is seen on a separate piece of rock. Length and 
height subequal; infero-proximal angle a right angle, upper margin 
convex. Outer surface slightly convex from end to end, more so from 
top to bottom. Inner surface equally convex in both directions, with 
a furrow close to the proximal end ; an oblique furrow near the lower 
edge, directed slightly upward distally, and a depression leading to 
the fdnus between the fingers and to the adjacent part of the immov- 
able finger ; on the oblique raised line below this depression, there is 
a row of 4 granules. Upper and lower edges margined; the lower 
edge shows, in the impression, about 18 dots, but whether these are 
punctae or spinules on the outer surface can not be told. There is a 
line of 6 punctae just' within the upper margin. A few scattered 
punctae on inner surface. An oblique cross section of the thumb is 
small and somewhat diamond-shaped; above this section there is a 
blimt tooth ; sinus between the 2 fingers V-shaped. The impression 
shows a little more of the length of the thumb than remains in the 
type, but represents neither the full length nor width. 

Paratype (a). — Left manus, larger than the holotype, and free 
from the matrix; immovable finger broken off; edges worn; inter- 
digital tooth present, and near by on the outer surface, a granule. 

Paratype (&). — ^A fragment, comprising the distal lower end of 
the propodus with broken finger attached ; interdigital tooth present. 



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148 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

CALLIANASSA HILU, mw iimcIm. 

IMute 58, figs. 18>20. 

Type-locality, — Gatun beds, Panama Canal Zone. Gkttun forma- 
tion. Miocene series. Robert T. Hill, collector; Station 18. 

Types,— C2Li. No. 135218, U.S.N.M. 

MeasureTnents. — Greatest height of carpus of right cheliped, lO^ 
mm. ; superior length of same, 10.3 mm. 

Material. — 3 pieces of rock from the same place contain fragments 
of Callianassa which belonged to at least 2 specimens and probably 
represent a single species. The largest piece contains 2 fragments, 
each consisting of a right merus and carpus (the better preserved of 
these may be considered the holotype) ; the imprint of a right wrist 
and hand; the distal half of a right propodus and a portion of the 
dactylus. The second piece of rock has been broken from the first 
and contains the continuation of the right propodus, with carpus 
and merus attached (paratype a), also the first 4 segments of the 
third right leg. The third piece of rock contains the sixth joint of the 
tail (paratype &). The following description is compiled from all 
the specimens: 

The merus of the right cheliped has the outer margin prominently 
carinate with a smooth rounded carina; upper margin granulated. 
In both instances the merus is so flexed beneath the carpus that the 
lower margin is not visible. The carpus is about equally long and 
high ; its upper margin is slightly convex to a point near the articula- 
tion with the merus ; distal margin slightly concave, but nearly ver- 
tical ; margin from the infero-distal angle to the merus strongly arcu- 
ate. Manus about as long as high, moderately convex, outer surface 
more so than inner; lower margin nearly straight; just above it on 
inner surface a row of many fine punctae from which setae may have 
sprung; this row is continued on the propodal finger; the latter is 
only partly uncovered ; it is flat on the inner side, at least half as long 
as the manus and its lower margin is a straight line continuous with 
that of the manus. 

The right leg of the third pair is very much like the corresponding 
member in C. stimpsoni Smith, the CaJManassa of the Atlantic coast 
of the United States. 

The sixth segment of abdomen or tail is subrhomboidal, with a con- 
striction behind the middle; the segment is much wider in front 
than behind ; the depressed portion at the anterior middle was hidden 
in life under the fifth segment. 



CALLIANASSA VAUGHANI. new i 

Plate 63, figs. 10-13. 
Type-locality. — Panama Canal Zone. From 85-foot cut on north 
side of big swamp on relocated line, Panama Railroad ; 1| to 2 miles 



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GBOLOOT AND PALEONTOLOGY OP THE CANAL ZONE. 149 

beyond Camp Cotton toward Monte Lino. Gatun formation. 
Miocene series. D. F. MacDonald and T. W. Vaughan, collectors. 
1911. Station 6030. Holotype and 2 paratypes. Cat. No. 324285, 
U.S.N.M. 

Holotype. — ^Portion of the right claw, comprising the distal end of 
the manus and both fingers; outer surface only. Height of manus at 
distal end equal to length of dactylus measured on the chord from the 
tip to the middle of its articulating edge; surface convex from upper 
to lower edge. Fixed finger wide in its proximal half, then abruptly 
narrowed; distal half directed slightly upward; length twice as great 
as basal width. The ends of the fingers are somewhat crushed ; the 
movable finger curves strongly downward and appears to overreach 
the tip of the immovable finger, being at right angles to it; its lower 
edge has two teeth, one near the articulation squarish, broader than 
long, the other smaller, separated by a rounded sinus. At the sinus 
distal to this tooth is the widest interdigital gape ; a little further on 
the fingers would meet for a ways, if they were closed, while tiie tips 
would cross each other. The greatest width of the dactylus is a little 
more than a third of its greatest length, measured in a straight line. 

The oblique edge of the propodus which projects over the dactylus 
is crenulate. On the manus near the gape of the fingers there are 3 
tubercles in a curve parallel with the edge. Behind the crenulation 
there are 2 scale-like sockets from which setae probably arose, and 
behind these a vertical thumb-nail impression. Near the supero- 
distal angle of the manus there is another socket. On the proximal 
half of the dactylus there are 9 sockets of larger size than those on 
the manus and irregularly disposed ; 2 are on the larger prehensile 
tooth and one on the smaller. 

Paratyfe {a). — Left manus, both sides visible. A smaller speci- 
men than the holotype. Upper and lower margins distally converg- 
ing. Length a little more than greatest width. Outer surface con- 
vex in both directions, but more so from top to bottom. Surface for 
the most part smooth and shining. The ornamentation is like that 
of the type, that is, a crenulated edge on the lobe which overlaps the 
dactylus, a row of tubercles just behind the gape — ^the lower of the 
3 tubercles is broken off — above this 2 sockets, and then a very short 
thumb-nail impression, followed by a socket near the upper angle. 
The Tipper margin is subacute in its proximal half, becoming gradu- 
ally blunt toward the distal end ; on either side is a row of sockets ; 
3 are visible on the outer surface and 5 on the inner; lower margin 
sharp, also with a row of sockets on either side ; 5 are visible on the 
outer side and about 10 on the inner side in the distal half; the 
proximal half is broken. There are a few punctae scattered about 
the outer surface; while on the inner surface near the depression 



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150 BULLETIN 103^ UKITBD STATES NATIONAL MUSEUM. 

leading to the gape there is a double row of granules, 8 of which 
can be made out. The edge overlapping the dactylus is crenulate on 
the inner surface. 

Paratype (&). — Portion of left propodus showing part of finger 
and a small part of the manus. Surface shining. Three tubercles 
near gape, and continuing the same curve along the upper part of the 
outer surface of the finger, there is a granule followed by 2 sockets. 

Mea8wrement8. — (Approximate only.) Holotype: Distal height 
of manus, 19.3 mm. ; length of dactylus from tip to proximal end of 
upper margin, 23.5 mm.; height of dactylus, measured straight up 
from the edge of the basal tooth, 8.6 mm.; length of immovable 
finger measured along the prehensile edge, 15 mm. ; height of same 
at base, 7 mm. Paratype (a) : Length of manus across middle, 19.5 
mm.; proximal height, 16.7 mm.; distal height, 15 mm.; greatest 
thickness, 6.6 mm. Paratype (b) : Height of immovable finger at 
base, 5.8 mm. 

Additional localities. — Panama Canal Zone. Las Cascadas sec- 
tion. From lower part of lime-cemented soft gray to olive -colored 
limestone with central parting of dark clay. The first hard, limy 
sandstone bed above the lower limestone just above Station 6019&. 
Upper part of Culebra formation. Oligocene series. D. F. Mac- 
Donald and T. W. Vaughan, collectors. 1911. Station 6019c. One 
left propodus with most of the fixed finger attached. The specimen 
is so bruised and crushed that its identity can not be determined with 
certainty. Cat. No. 324283, U.S.N.M. 

Also, from the same place, a right dactylus from a cheliped of 
much smaller size. Its identity is uncertain. It lacks the large teeth 
on the cutting 6dge, but it may belong to the feebler of the two 
chelipeds, or to a female. There is evidently a shallow sinus at the 
base, followed by a low broad tooth. Six sockets for setae can be 
made out. The thick outer crust has nearly all broken away. Cat. 
No. 324283, U.S.N.M. 

Panama Canal Zone. From lowest horizon in big cut from i to -J 
mile beyond Camp Cotton toward Monte Lirio. Lower part of 
Gatun formation. Miocene series. D. F. MacDonald and T. W. 
Vaughan, collectors. 1911. Station 6029a. Left manus, about 9.6 
mm. long, measured at the level of the articulating condyle of the 
dactylus; edges mostly broken and obscured. Identification based 
(1) on the general contour of the surface, (2) the color, a light drab, 
(3) the margin adjacent to the articulating condyle of the dactylus, 
and (4) 2 sockets just below the upper margin on the inner surface. 
Cat. No. 324284, U.S.N.M. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 151 

CALUANASSA 8TRIDEN8, new spMieik 

Plate 61, figs. 12-14. 

Type-locality, — Panama Canal Zone. From third hard sandstone 
bed from bottom. Las Cascadas section. Upper part of Culebra for- 
mation. Oligocene series. D. F. MacDonald and T. W. Vaughan, 
collectors. 1911. Station 6019^. Cat. No. 324281, U.S.N.M. 

Holotype. — Manus of a left cheliped. Only the upper two-thirds 
is visible, the lower third is embedded in rock. Upper margin 
horizontal, distal and proximal margins vertical, supero-posterior 
comer rounded. Upper edge thin, a little sinuous, viewed from the 
top. On the inner surface a little below the upper edge there is a 
row of 8 short vertical ridges, which occupies the whole length of 
the segment. This may have been a stridulating mechanism. 

Measurements. — Superior length of manus, 11 mm.; thickness, 
3.1 ram. 

CALLIANASSA MAGNA, new ip^ieik 

Plate 62, figs. 1-3. 

Type-locality. — ^Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. From lowest fossiliferous bed. Third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
lialf of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a. Cat. No. 
324270, U.S.N.M. 

Measurements. — Greatest length of movable finger (tip broken 
off), 33 nun.; greatest height, 13.2 mm.; thickness, 0.8 mm.; length 
of basal sinus, 6 mm. 

Holotype. — ^The only specimen is a movable finger or dactylus of 
the right cheliped. It is very much worn and a considerable portion 
of the tip is lacking. The lower border is thin, but the remainder 
is thick and in cross section subcircular, and tapers gradually to- 
ward the distal end. At the proximal end below there is a broad 
sinus; the thin prehensile edge is slightly concave, viewed from out- 
side, and feebly denticulate, especially when viewed from inside; 
there is a somewhat larger and better preserved tooth just within the 
margin at the widest part of the finger. Upper margin in outer view 
straight in its proximal half, gently curved distally. In dorsal 
view the finger is much curved and in the middle of its upper surface 
there is a longitudinal row of four large punctae. 

In its general shape, including the basal sinus, this dactylus re- 
sembles that of C. pellucida Rathbun, from the Leeward Islands, 
a description of which is about to be published by the Carnegie In- 
stitution, but the prehensile edge is thinner and more laminate and 
the inner outline more concave in dorsal view instead of almost 
f^traight as in that species. 



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162 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



CALLIANA88A CBA8a4, b«w 

Plate 61, flgt. 1-8. 

Type-locdUty. — Gatun section A, bed A (bottom of section). 
Lower part of Gatun formation. Miocene series. D. F. Mac* 
Donald, collector. Station 6003. Two dactyli of the left cheliped- 

Types.— CiLt. No. 324276, U.S.N.M. 

Measurements. — ^Length of dactylus, 16.3 mm. ; height, 6.2 mm. 

Holotype. — Outer and upper surfaces exposed, outline of tip ob- 
scure. Viewed from the outside the upper outline is arcuate and the 
tip bent down below the prehensile edge, which is nearly horizontaL 
Viewed from above, the outer line is much curved and the inner line 
nearly straight. On the prehensile edge there is a shallow basal 
sinus, followed by a very low, broad tooth; rest of margin faintly 
sinuous. The surface shows a number of granules, some large, others 
small: the large ones are about 7 on the outer surface and 5 on the 
upper surface; of the former, 2 are submarginal, one of them being 
above the lobe, the other half way to the tip; the other 5 external 
granules are disposed on the distal half; the 5 superior granules are 
arranged in 2 rows, one row of 3 toward the inner surface, and the 
other row of 2 granules toward the outer surface; the proximal of 
these is double. On tlie middle of the outer surface there is a patch 
of about 50 small granules. The chalky-\\ hite outer layer of the shell 
has crumbled away except near the edges, so that one can not tell 
whether the granules were apparent on that surface. Color of sur- 
face now exposed dull light bluish. 

Paratype, — About t>vo-thirds as large as the holotype, and with 
both ends of the finger missing. Granules as follows : 3 large above 
the lower margin, 2 as in the holotype, the other above the basal 
sinus; on the inner surface are 3 similarly spaced but placed more 
distad. On the upper surface there is a row of 4 toward the inner 
surface, and below the second one from the proximal end are 2 near 
iogether. The small granules are more separated than on the holo- 
type and are distributed chiefly on the upper half of the outer 
surface. 

CALLIANASSA. spedM. 

Plfito 59, fig. 5. 

Locality, — Panama Canal Zone. Top part of limy sandstone be- 
low upper conglomerate, near foot of stairs, Gail lard Cut. Upper 
part of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6012r. Cat. No. 
324277, U.S.N.M. 

Material. — ^Manus of left cheliped of a small specimen. Outer sur- 
face visible. Very convex from top to bottom, a deep groove next to 
the edge articulating with the carpus; upper and lower margins ill- 
defined. 



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OBOLOGT AND PALEONTOLOGY OF THE CANAL ZONE. 153 

CALUANASaA ?, spmIm. 

Plate 64, fig. 10. 

Locality. — Panama Canal Zone. Las CascadaB section. From 
lower part of lime-cemented soft gray to olive-colored limestone 
(with central parting of dark clay). The first hard, limy sandstone 
bed above the lower limestone just above station 60196. Upper part 
of Culebra formation. Oligocene series. D. F. MacDonald and 
T. W. Vaughan, collectors. 1911. Station %019c. Cat No. 324280, 
U.S.N.M. 

Material, — One small specimen resembling in shape the merus 
joint of the smaller of the chelipeds of the first pair. If the above 
^^ess be correct, this is from the right cheliped. The two oblique 
grooves near the distal end may have been artificially produced. 

Family PAGURIDAE. 

PBTROCmBUS BOUYIERI. new gpedes. 

'* Petrochirus cf. granulatua Olivier sp,'* Toula, Jahrb. der k. k. Gteolog. 
Reichsanstalt, Wien, vol. 61, 1911, p. 511 [25], pi. 30 [1], fig. 13. 

Gatun; Miocene (Toula). Not represented in the collection at 
hand. 

There are two recent species of Fetrochirus on opposite sides of 
the continent, namely, P. hahamensis (Herbst)^=P. granulatua 
(Olivier), which extends from Florida to Brazil, and P. calif omi- 
ensis Bouvier^ taken at La Paz, Mexico, and in Ecuador. One of 
the principal differences between them lies in the ornamentation of 
the chelae. The right chela of P. hahamensia is covered chiefly with 
fan-shaped clusters of granules, all of which trend forward and 
present a smooth, oval side- face when viewed dorsally; the clusters 
vary in size, and some are composed of only 2 granules, while others 
are represented by only one granule ; all are fringed anteriorly with 
hair, which fills the interspaces. The right chela of P. califormensis 
has similarly clusters and single granules, but the clusters are not 
fan-shaped but round, or nearly round, and are composed of a large 
smooth central granule surrounded by small granules tipped with a 
sharp, homy point; the granules are much more elevated and have a 
more dorsal inclination than in bahamensis; the single granules are 
also more numerous than in that species. 

The right chela of the fossil specimen figured by Toula resembles 
that of P. caUfomiensis. 

The left chela of P. bahamensis is covered with fan-shaped clusters 
of granules like those on its right chela, but the clusters are more 

^ Cancer iHihamen^ii Herbst, Natorg. d. Krabben u. Krebse, toI. 2, 1796, p. SO. 
'Bull. Mos. mat. Nat, Paris, 1896, p. 0. 



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154 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

crowded, and single granules are fewer. The left chela of P, cali- 
forrdensiB is covered with clusters like those on its right chela but 
more crowded, and the granules of which they are composed are of 
a smaller average size. 

The left chela of the fossil agrees more nearly with that of P, 
bahamensis. 

We therefore have a Tertiary species combining the characters of 
two Recent species, at least as far as the chelae are concerned, possess- 
ing the right chela of one and the left chela of the other. 

Tribe BRACHYURA. 

Subtribe Dromiacea. 

Fanuly DROMnDAE. 
Genus 60NI0CHELE BelL 

GONIOCHBLE? ABMATA, b«w ■ptriM. 

Plate 57. figs. 11 and 12. 

Type-locality. — Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. From lowest fossiliferous bed. Third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a, Movable 
finger of the left cheliped. Cat. No. 324259, U.S.N.M. 

Measurements. — ^Length of movable finger (tip broken off) , 19 mm. ; 
width at about the middle, 6 mm. ; greatest thickness, 4.5 mm. 

Holotype. — ^The shape is elongate-triangular viewed from outside, 
the prehensile edge being nearly straight and the upper edge slightly 
arched; outer surface convex in both directions. The prehensile 
edge has a sinus at its base, defined distally by a broad tooth which 
is at present truncate, but may have been prolonged in two small 
teeth ; beyond are four teeth separated by rounded sinuses and with 
their tips missing; the first two are spiniform, the last two broad 
and thin. The upper margin bears 5 low, spaced teeth, while just 
within and alternating there is a line of 3 teeth. On the outer sur- 
face are 2 rows of tubercles not far from the margins, 4 in the lower 
and 3 in the upper row; the proximal tubercle in the upper row 
is bifid. On the inner surface are 5 tubercles besides those above 
mentioned, 2 in a longitudinal row in the middle, 2 on the distal half 
a little above the prehensile edge, and 1 small one toward the top 
and behind the middle. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 166 

I have placed this specimen in Goniochele^ on account of the 
dactylus being armed on both edges, as in G^. angvlata Bell,* the 
type of the genus, and because the general shape of the segment is 
similar. In Bell's species the dorsal surface is smooth. 

Subtribe OXYSTOMATA. 
Family CALAPPIDAE. 

HEPATUS CmLIENSIS Bliliie Edwardi. 

Plate 66, fig. 4. 
Hepatus chiliensis Milne Edwards, Hist. Nat. Crust., vol. 2, 1837, p. 117. 

Locality ' — ^Panama Canal Zone. From near Mount Hope in ditch 
through swampy ground. About one-quarter mile from present sea 
beach, 6 to 8 feet above high tide. Pleistocene series. D, F. Mac- 
Donald, collector. April, 1911. Station 5850. Cat. No. 324235, 
U.S.N.M. 

Material. — Dactylus of right chela, 5.3 mm. long. This little 
specimen shows all the essential characters of recent individuals of 
this species : 8 shallow rounded teeth on the lower margin ; a row of 
5 tubercles on the proximal part of the upper margin and a row of 
4 tubercles just below and on the outer surface; still further down, 
2 more tubercles; a stridulating ridge on the inner surface just 
below the upper edge is formed of upwards of 45 fine parallel striae 
and occupies the greater part of the length of the finger. 

Distribution of Recent material. — ^Eanges at the present time from 
Ecuador to Chile. 

HEPATUS, BpeciM. 
Plate 66, fi^. 12. 

Locality. — Panama Canal Zone. From lower part of lime-cemented 
<?oft gray to olive-colored sandstone (with central parting of dark 
clay)* The first hard limy sandstone bed above the lower lime- 
stone just above fossil lot No. 6019&. Upper part of Culebra forma- 
tion. Oligocene series. D. F. MacDonald and T. W. Vaughan, 
collectors, 1911. Station 6019^. Cat. No. 324239, U.S.N.M. 

Material. — One dactylus of left chela, much worn and incom- 
plete at both extremities; the proximal half of the upper margin is 
also wanting. The curves in side view are much like those of 
H. cMliensis Milne Edwards.* Both inner and outer surfaces are 

> A Monograph of tbe Fossil Malacostracoas Crustacea of Great Britain, pt. 1, 1807, 
p. 25. 

* Idem, pi. 4, fig. 0. 

•HlBt. Nat. Crust., vol. 2, 1837, p. 117. 

8370'--18— BuU. 103 11 



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166 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

convex from top to bottom. The prehensile edge has traces of 2 
small teeth, one near the base and one at about the distal third. 
There are also various depressed granules or pits, namely, a row of 
5 small ones on the upper margin; opposite the distal one of these 
there is a larger one on the inner surface; and 3 or 4 on the outer 
surface. These are all sunken so that they do not project above the 
present surface, which is not the true outer surface. 

This jBnger is not quite so flat inside as H. cfiiliensis. 

Measurements. — Length of dactylus, 13.6 mm.; height at middle^ 
3.6 mm. ; thickness at middle, 2.7 mm. 

CALAPPA COSTARICANA. new specks. 

Plate 57, fig. 24. 

Type-locality. — Costa Rica: City of Port Limon. Port Limon 
formation. Pliocene series. Dr. L. A. Wailes. 4269. 

n olotype,— C^it Xo. 324240, U.S.N.M. A triangular fragment 
measuring about 9 mm. on each margin, representing the propodal 
finger and the infero-distal portion of the palm of a left chela of 
the weaker form — ^that is, without a strong submarginal tooth or lobe 
characteristic of the stronger chela in Calappa. Lower margin very 
sinuous, the tip of the finger directed upward, proximal half of 
margin armed with 6 strong tubercles directed distally. Just above, 
on the outer side, and beginning nearer the finger-tip there is a row 
of 11 smaller tubercles, normal to the surface. The prehensile edge 
is nearly straight, inclined at an angle of about 75° with the lower 
margin of the palm, and is furnished with 6 large, unequal tubercles, 
which end at the small sinus which ordinarily exists just below the 
raised margin surrounding the articulation with the dactylus; on 
the upper side of this sinus are 2 small tubercles, and above this 
point, the specimen comes to an end. The outer surface is covered 
with upward of 40 flattened scale-like tubercles pointing upward; 
they are separated from the submarginal row by a smooth depres- 
sion. The propodus is thick and the inner surface is beveled, the 
bevel for the most part smooth ; remainder of inner surface studded 
with very unequal pearly granules and tubercles; interspace crowded 
with fine punctae ; 2 sinuous ridges run toward the finger-tip. 

This species resembles C. gallus (Herbst)^, which is fomid living 
at the present time from Florida Keys to Bahia, Brazil. The fossil 
species differs in the very prominent tubercles of the lower margin 
of the palm and the longer tubercles of the submarginal row just 
above, on the outer surface. 

* Cancer gallus Herbst, Natur. Krabben u. Krebse, vol. 8, pt. 8, 1808, pp. 18 and 46, 
pi. 68, fig. 1. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 157 

CALAPPA FLAMMEA (H«rlMt). 

Plate 61. figs. 4 and 5. 

Cancer flammeus Herbst, Naturg. d. Krabben u. Krebse, vol. 2, 1794, p. 
161, pi. 40, fig. 2 ; vol. 3, pt. 3, 1803, p. 19. 

Locality. — Near Mount Hope, Panama Canal Zone, in ditch 
through swampy ground about one-fourth mile from present sea 
beach, 6 to 8 feet above high tide; Pleistocene series; D. F. Mac- 
Donald, collector. April, 1911. Station 5850. Cat No. 324237, 
XJ.S.N.M. 

Represented only by one dactylus or movable finger belonging to 
the stronger chela. The milling of the stridulating ridge on the 
inner surface just below the upper edge is more strongly ma'rked than 
in most of the recent specimens examined. 

MecLmremerUs. — ^Extreme length, 15 mm.; width just distal to the 
upper marginal tooth, 4.7 mm. 

Distribution of Recent material. — From North Carolina to Colom- 
bia and Venezuela. 

CALAPPA ZURCHBRI Bovrter. 

Calappa zurchen Bouviek, Bull: Mus. Hist. Nat. Paris, vol. 5, 1899, p. 190, 
text-fig. 

Panama. Lower Miocene. 

Not represented in the Museum collection. 

CALAPPELLA, new genus. 

Carapace very little broader than long, without clypeiform ex- 
pansions, but with a spine at the junction of the antero-lateral and 
postero-lateral borders, and a spine at each end of the posterior 
border. 

Front small, projecting forward beyond the orbits. 

Orbits small, directed forward. 

In the narrow front and small orbits, this genus resembles Calappa^ 
but in its narrow carapace armed with 4 slender spines, it differs 
from that genus as well as from all other Calappinae. 

Type of the genus. — Calappella quadrispina^ new species. 

CALAPPELIA QUADRI8PINA, new spmIm. 

Plate 58, figs. 1 and 2. 

Type-locality. — ^Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut From lowest fossiliferous bed; third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 



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168 BUUJETIN 103, UNITED STATES NATIONAL MUSEUM. 

and T. W. Vaughan, collectors. 1911. Station 6020a. Cat. No. 
824288, U.S.N.M. 

Measurements (approx.). — Length of carapace (spines excluded), 
14 mm.; width (spines excluded), 14.4 mm. 

Holotype. — Species represented by one specimen showing part of 
the carapace and no appendages, enclosed in a nodule which has been 
broken in two. Nodule not much larger around than the carapace. 

Carapace nearly as long as wide, spines excluded; width between 
outer angles of orbits a little greater than posterior margin. Chord 
of the antero-lateral margin nearly twice as long as that of the 
postero-lateral margin. Anterolateral margin divided into 2 parts, 
the anterior two-fifths being slightly convex, the posterior three- 
fifths very convex with a tubercle at its middle. Postero-lateral mar- 
gin somewhat sinuous, but in general concave; posterior margin 
slightly convex, about twice as long as the slender spine at either end 
which is directed backward and slightly outward. Lateral spine also 
slender, but longer, half as long as the postero-lateral margin and 
pointing obliquely backward. 

The central and anterior part of the surface of the carapace is 
lacking. There are, however, two oblique, parallel, branchial fur- 
rows; between them a row of 8 tubercles and some scattered granules; 
the outer part of the branchial region is higher and rough with 
irregular tubercles which are more or less confluent. Cardiac region 
high, with a median tubercle on its posterior slope; in almost the 
same plane transversely, but on a lower level, there is another tubercle 
on each side just above the postero-lateral margin. 

There is a very small hollow in the nodule where the point of the 
front rested, and on the other half of the nodule may be seen the 
lower surface of the front where it joined the interantennular septum. 

The orbits are small, their upper and lower margins formed by 
two teeth, the innermost advanced, separated by a blimt V-shaped 
sinus. 

On the lower surface, the inner tooth of the orbit is considerably 
elevated (that is, ventrally). A sharp ridge runs obliquely back- 
ward from or near the epistome, and is armed with a tooth at its 
posterior third. 

MUBSIA MACDONALDI, new spcdM. 

Plate 58, fig. 21. 

Type-locality. — Panama Canal Zone. Las Cascadas section. Fifth 
or topmost limestone. Emperador limestone. Oligocene series. D. 
y. MacDonald and T. W. Vaughan, collectors. 1911. One specimen, 
part of left cheliped. Station eO\9g. Cat. No. 824229, U.S.N.M. 



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GEOLOGY AKD PALEONTOLOGY OF THE CANAL ZONE. 159 

Measwrementa. — ^Lieiigth of palm between articulations, 19 mm.; 
height (approx.), 10.2 mm. 

Represented by the left palm only, which has the customary Calap- 
poid form, and a portion of the immovable finger; the edges are not 
well shown ; two teeth may be seen near the distal end of the upper 
margin. The only details of the outer surface that can be made out 
are a few large tubercles, more or less compressed as in Kecent species 
of MuT^; these tubercles number about 15; there is a row of 4 
a little above, and subparallel to, the lower margin; from these 4 
tubercles irregular rows extend obliquely upward, trending toward 
the fingers; including those of the horizontal row, those of the distal 
oblique row are 4, of the second row 5, of the third row 3, of the 
proximal row 3 ; these rows are not strictly parallel nor their tuber- 
cles regularly spaced. The lower proximal tubercle is the largest and 
most compressed. 

There is an indication of the distal spine of the arm- joint, which 
may be seen in the figure. 

Eesembles Mursia armata de Haan,^ but in that species the prin- 
cipal tubercles of the hand are 9, arranged in 3 parallel and fairly 
regular rows. 

MURSIA OBSCURA, new species. 
Plate 61, fig. 18. 

Type-locality. — Near Panama Canal Station "1910," north of 
Pedro Miguel locks, Panama Canal Zone. From dark clay, lower 
part of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6010. Cat. No. 
324226, U.S.N.M. 

Measurements of fragmentary specimen, 8.2 mm. long, 9 mm. wide. 

Holotype. — The central part of the carapace, devoid of its margin, 
except perhaps the middle of the posterior margin. This fragment 
is embedded wrong side up in a piece of rock; only the thin outer 
crust of the specimen remains and its under surface alone is visible! 
It has been referred to this genus because the cavities or pits, which 
represent tubercles on the dorsal surface of the shell, are arranged 
much as in Mv/rsia. There are 5 longitudinal rows of these pits : The 
median row consists of 3 large pits., one cardiac, one genital, one gas- 
tric, preceded by 2 small pits side by side; the two lateral rows 
(on each side) are very little oblique to the median line, but sub- 
parallel to each other; the inner of these rows consists of 4 pits, the 
penultimate one being in a transverse line with the last pit of the 
median row ; the last pit of the inner lateral row is round and deep 

1 Fauna Japonic^, 1880» p. 78, pi. 19, fig. 2. 



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160 BULLETIN 103, UNITED STAIBS NATIONAL MUSEUM. 

and indicates a large excrescence, perhaps a spine, on the dorsal sur- 
face; 3 pits only are visible in the outer row, the middle pit is in 
transverse line with the anterior pit of the other rows, while the 
anterior pit of the outer row is not quite in line with the two behind 
it. Either side of the pair of small submedian gastric pits there is 
a large cavity. The furrows separating the branchial from the 
gastric and cardiac regions are indicated by sinuous ridges. 

Judging from the proximity of the rows of pits, this carapace is 
narrower in proportion to its length than in other species of Murda. 
This together with the evidence of a strong prominence near the 
postero-lateral borders points to a genus different from any described. 

MURSILIA, new senus. 

The manus resembles that of Mursia, but lacks the crest or ridge 
on the inferior margin. 

Type of the germs. — MursiUa ecriatata^ new species. 

MURSILIA ECRI8TATA, new ipedM. 

Plate 57, fig. 27. 

Type-locality. — Gatun beds. Gatun formation. Miocene series. 
"Robert T. Hill, collector. 

Holotype.—C^A. No. 135219, U.S.N.M. 

Measurements. — Length of palm, 9.8 mm., height of palm, 7.4 mm. 

Represented by only one specimen showing the right palm and a 
portion of the wrist. Palm short and high. Surface finely and 
rather distantly granulated on the upper half of the outer surface 
and at the proximal end ; more closely granulated on the lower sur- 
face. There are 9 large tubercles arranged in 3 oblique, subparallel 
rows ; the tubercle at the inferior proximal comer is much the largest, 
is flattened above and has a raised rim; between it and the next 
tubercle in the horizontal row, but a little below, there is a smaller 
tubercle. Below the distal tubercle of the horizontal row of 3, and 
nearer the inner than the outer surface there is a small tubercle. On 
the upper margin there are 7 narrow, thickened teeth similar to those 
of Calappa. Below the sinus between the fifth and sixth teeth 
(counting from the wrist) there is a low tubercle; also one on tire 
base of the second tooth. A part of a tubercle near the beginning of 
the immovable finger is visible. 

The outline of the wrist is defined, but very little of the surface 
remains; a small piece near the distal upper comer is granulated like 
the upper half of the palm. 

The tuberculation of the manus or palm resembles that of Mursia^ 
the dentation of the upper margin is nearer that of Calappa, while 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 161 

-the segment differs from both those genera in lacking the crest on the 
lower margin of the pahn. 

Family LEUCOSHDAE. 

LEUCOSILIA JURINEI (Saiusare). 

Owiia (ilia) jurinei Saussube, Rev. et Mag. de Zool., No. 8, 1853, p. 12, 

pL 13, fig. 4. 
Leucosilia jurinii Bell, Trans. Linn, Soc. Ix»ndon, vol. 21, 1855, p. 295, pL 82, 

fig. 1. 

Locality. — Panama Canal Zone. From near Moimt Hope in ditch 
through swampy ground. About one-quarter mile from present sea 
beach, 6 to 8 feet above high tide. Pleistocene series. D. F. Mac- 
Donald, collector. April, 1911. Station 5850. 

Material. — Six small arm-joints more or less worn; four of them 
iu*e incomplete at one or both ends. 

Siee. — ^Length of a large one, 5.6 mm. 

Distribution of Recent Material, — ^Ranges from Mazatlan, Mexico, 
to Peru and the Galapagos Islands. 

LEUCOSILLk BANANENSIS. new species. 
Plate 57, figs. 6-8. 

Type-locality. — ^Banana River, Costa Rica. Probably equivalent 
to Gatun formation. Miocene series. D. F. MacDonald, collec- 
tor. 1911. Station 5882A, 5&, 1 arm, holotype, from seventh fos- 
silif erous zone below the uppermost one of the section. Station 5882^, 
5a, 1 arm, paratype {a) , from sixth fossilif erous zone below the up- 
permost one of the section. Station 5882/, 3/, 1 arm, paratype (&), 
from fifth fossilif erous zone below the uppermost one of the section. 

ryp6«.— Cat. Nos. 324230, 324231, and 324232, U.S.N.M. 

Measurements. — ^Length of holotype, 11.1 mm. ; greatest diameter, 
4.2 mm. 

Bepresented by only 3 arms from 3 different layers. The best 
specimen represents the left arm nearly complete, lacking only the 
•distal articulating edge. 

Shape subcylindrical, slightly compressed in a vertical direction, 
the greatest diameter being proximal to the middle, the smallest 
diameter at the proximal end. The ornamentation consists of 
tubercles or large granules, the granules becoming small at both ends 
of the arm ; around the middle of the segment the granules number 
about 15 ; the tops of the granules are broken off so that they appear 
much flatter than they really were. Compared with L. jurinei^ the 
jirm is more swollen, the granules less numerous, more equal and 
further apart. 



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162 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

LEUCOSnDAE?, senns and species indeterminable. 

Plate 60, figs. 7 and 8. 

Locality — Panama Canal Zone. From near Mount Hope in ditch 
through swampy ground. About one-quarter mile from present sea 
beach, 6 to 8 feet above high tide. Pleistocene series. D. F. Mac- 
Donald, collector. April, 1911. Station 5850. Cat. No. 324236, 
U.S.N.M. 

Material. — ^Dactylus of left chela, 11.4 mm. long, with proximal 
end lacking. This dactylus differs from those of Persepkona and 
allied genera in its strong curvature upwards, supposing the pre- 
hensile edge to be directed inwards. This edge is nearly straight 
except just at the tip and is armed with 25 small unequal teeth, 2 of 
which in the proximal third are the largest. Besides this edge the 
surface is composed of 4 high, smoothly rounded ridges separated 
by narrow grooves ; 2 of the ridges are inferior, and 2 superior, the 
outermost of the latter embracing the outer edge and having a longi- 
tudinal row of punctae near its middle. Each side of the prehensile 
edge there are 2 or 3 rows of punctae. Tip of finger bent rather 
abruptly but obliquely inward, while in its upward trend it continues 
the curve of the rest of the dactylus. 

Subtribe BRACHYGNATHA. 

Superfamily BRACHYRHYNCHA. 

Family PORTUNIDAE. 

CALLINECTES DECLIVIS, new tpeclM. 
Plate 66, figs. 1-3. 

Type-locality. — Banana Eiver, Costa Rica. Eighth fossiliferous 
zone below the uppermost one of the section. Probably equivalent 
to Gatum formation. Miocene series. D. F. MacDonald, collector. 
1911. Station 5882^ 6c. Cat. No. 324262, U.S.N.M. 

Measurements. — Greatest height, 14 mm.; length of manus meas- 
ured horizontally from extreme base of proximal spine, 21 mm.; 
thickness, 9.2 mm. 

Holotype. — ^The propodus of the left cheliped, with the tip of the 
finger broken off. The palm is prismatic as in recent species of 
CaZlinectes^ with 7 facets, more or less distinct ; 4 facets on the outer 
surface and 3 on the inner surface. The surface, or what remains of 
it, is smooth and shining to the naked eye, but under a lens, shows 
very fine granulation, and larger scattered punctae. The facets are 
separated by blunt ridges; one facet is a little above the middle of 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 163 

the outer surface, and has subparallel margins; the facet below this 
widens distally and its lower edge, not very prominent, is continued 
upon the finger, where it is sharper; the lower facet of the outer sur- 
face is not sharply marked; the facet above the middle narrows 
slightly toward eitiier end and has raised margins; at its proximal 
end there is the stout base of a large spine such as exists in CalUnectes 
of the present day. The upper facet of the inner surface is narrow, 
wider in the middle than at the ends, and is not depressed, its distal 
outer corner only is visible when the manus is viewed externally; 
at the distal end just outside the inner margin is an indication that 
a spine has been broken off. The remainder of the inner surface is 
divided into 2 facets of nearly equal width separated by a prominent 
ridge. 

The finger is a little curved inward; on the outer and inner sur- 
face there are 2 grooves, each with a row of large punctae, the groove 
at the middle of each surface being deeper than that near the pre- 
hensile teeth. The teeth are of moderate size, irregular, the larger 
(mes alternating with one or two smaller ones; at the broad proximal 
end of the cutting edge there are 2 small teeth side by side, one near 
the inner the other near the outer surface. 

This propodus differs from those of all the Recent OaUinectes in 
the position of the uppermost facet. In (7. sapidus^ etc., this facet is 
a part of the outer series, that is, continues the slope of the adjoining 
facet on the outer surface; while in the fossil it inclines downward 
toward the inner surface except at the distal epd where it is nearly 
horizontal. Furthermore, the propodus is shorter in proportion to 
its height than in recent CdUinecte^. 

I have placed this species in CaUinectes rather than in Portwnus 
{^Neptunus of authors) because the palm is nearer the shape of 
Callinectes than it is to similar segments in the genus Portunus^ as 
P. aangmnolentus ; the fossil is very unlike any Portunus now living 
on the coast of tropical America. 

CALUNECTES RETICULATUS, new spMiei. 

Plate 66, figs. 5-7. 

Type-locality. — Panama Canal Zone. Las Cascadas section, Gail- 
Jard Cut. From lowest fossiliferous bed; third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a. Cat. No. 
324261, U.S.N.M. 

Measurements. — Greatest height of manus, 15.6 mm.; length of 
manus measured horizontally on middle of outer surface, 19.2 mm. ; 
thickness, 10 mm. 



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164 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Holotype. — ^The propodus of the right cheliped, the immovable 
finger being broken off near its middle. 

In shape, this hand is shorter, higher, and thicker than the pre- 
ceding. The surface, aside from the ridges and the uppermost facet 
is covered with a fine reticulation of transverse grooves. The facets 
are 7 in number and in position are like those of C. declivis^ except- 
ing that the narrow uppermost one which appears to belong to the 
inner surface in C. decUvis is more horizontal in C. reticvlatua and 
forms the upper surface of the segment. The next facet on the outer 
side is narrowest at the distal end and widens to the middle, after 
which the margins are subparaUel; the next facet widens distally, 
and the next also, but in a lesser degree; the lower facet is ill defined. 
The 2 facets of the inner surface are subequal and widen distally. 

There is the base of a tooth at the inner distal end of the upper 
facet, and a short blunt spine at the distal end of the crest between 
the upper and middle facets. If there was a tooth at the proximal 
end it is broken off. 

The propodal finger bears on its upper edge the stumps of 3 sub- 
equal teeth ; nearer the palm on the same surface there are 2 small 
acute tubercles transversely placed, the inner one the larger. 

Aside from the difference in shape and ornamentation between this 
species and the preceding, there is a difference in the form of the 
facets which may be seen by comparing figures 5 to 7 with 1 to 3 on 
plate 66. 

CALLINECTES, ipacIm. 

Plate 65, fips. 1 and 2. 

Locality. — Panama Canal Zone. From the 4 feet of dark, strati- 
fied tuff and clay immediately overlying the lower limestone bed, 
Las Cascadas section. Upper part of Culebra formation. Oligo- 
cene series. D. F. MacDonald and T. W. Vaughan, collectors. 
1911. Station 6019&. Cat. No. 324255, U.S.N.M. 

A specimen of a left manus and carpus, very much worn, the upper 
and lower margins and the distal end of the manus being lacking. 
Tlie palm is narrower than in C. reticul<itus described above; the 
facet near the middle of the outer surface is wider than in C. decUvis^ 
and widens distally instead of having subparaUel margins as in that 
species. 

CALUNECTES, spMiet. 

Plate 65, flg. 7. 

Locality. — Panama Canal Zone. From top part of limy sandstone 
below upper conglomerate, near foot of stairs, Gaillard Cut. Upper 
part of Culebra formation. Oligocene series. D. F. MacDonald 



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QBOLOGY AND PAI£0NTOL0QT OF THE CANAL ZQNB. 165 

and T. W. Vaughan, collectors. 1911. Station 6012c. Cat. No. 
324268, U.S.N.M. 

The distal third of the immovable finger of a claw of a Callinectes. 
On the prehensile edge is shown the most distal of the large teeth 
customary in the genus followed (toward the tip) by 3 smaller teeth ; 
the tip is defective, having been broken off, then reattached in the 
wrong place. There is a punctated groove down the middle of the 
inner and the outer surface. 

Of the species of CaOmeotes living on the Pacific coast of America, 
this fragment resembles most C. toxotea Ordway,* which occurs from 
Cape St. Lucas to Peru. 

ARENAEUS, 



Plate 64, fig. 1. 

Locality. — Panama Canal 2k)ne. From near Mount Hope in ditch 
through swampy ground. About one-quarter mile from present sea 
beach, 6 to 8 feet above high tide. Pleistocene series. D. F. Mac- 
Donald, collector. April, 1911. Station 6850. Cat. No. 324252, 
U.S.N.M. 

Material. — Five fingers worn and more or less incomplete. Prob- 
ably all are movable fingers or dactyli. Length of most perfect 
specimen, 8.2 mm. On the outer surface there are two grooves dotted 
with minute puhctae; one is shallow and near the prehensile teeth, 
the other is above the middle of the segment; on the upper surface 
there are also two punctated grooves, but near together, while the 
inner surface has two furrows similar to those of the outer surface. 
Three or four of the prehensile teeth are enlarged as is usual in 
Portunids, and the tip is curved downward. There are evidences of 
close granulation on the uppermost ridges and on the proximal part 
of the segment. 

This is near A. mexicaims (Gerstaecker*), a Recent species which 
occurs from the west coast of Mexico to Peru. The shape, curvature, 
and granulation are similar, but three of the six grooves belong defi- 
nitely to the outer surface. 

EUPHTLAX CALLINECTIAS, new spMiet. 

Plate 65, figs. a-6. 

Type-locality. — Banana River, Costa Rica ; ninth f ossilif erous zone 
below the uppermost one of the section. Probably equivalent to Gatnn 
formation. Miocene series. D. F. MacDonald, collector. 1911. 
Station 5882;/ 5rf. Cat. No. 324234, U.S.N.M. 

^Boston Journ. Nat. Hist, vol. 7. 1863, p. 576. 

^Buotenoia mtricona Gcntaeckor, Arch. f(lr Naturg., vol. 22, pt. 1, 1866, p. 131, pi. 5, 
flga. 3 and 4. 



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166 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Medswernents. — ^Length of carapace, 45 mm. ; gastro-cardiac suture, 
9.6 mm. 

Holotype. — One male specimen, showing parts of the upper and 
lower surfaces, but no appendages. The outer layer of the shell of 
the carapace remains only in the central part, where the gastric, 
branchial and cardiac regions meet. This surface is granulated and 
areolated much as in CaUinectes;^ that is, the regions are separated 
by definite depressions, the gastro-cardiac groove is transverse, there 
is an areola at the inner angle of the branchial region, but there is a 
shallower division into 2 lobules than in CalUnectes. The post- 
gastric area is incomplete anteriorly, so that it is impossible to tell 
whether it is marked by a raised and granulated margin ; the shape 
of this area most nearly resembles that of C. exasperatus (Grer- 
staecker*), and its granulation that of C. hocourti A. Milne Edwards, 
the granules being absent or sparse along the lateral and posterior 
borders. Across the middle of the gastric region runs a blunt eleva- 
tion, concave forward. The branchial region is divided in two by 
a depression running obliquely backward and outward. These last 
two features suggest the carapace of Euphylax dovu Stimpson,* a 
species now existing on the Pacific coast of America, between Central 
America and Payta, Peru. 

Anterior margin very broad, as in E. dovii^ most of it being 
occupied by the orbits; the front is narrow, T-shaped, much con- 
stricted at base; the anterior part of the T has a concave surface 
and is deflexed to meet the epistomial spine. This is more advanced 
than the front; its tip is broken off. Tlie upper margin of <he 
orbit slopes backward and outward and is somewhat undulating. 
The shape of the orbit can not be definitely made out, but a portion 
of the smooth inner lining of the outer extremity remains. The 
indications are that the eyestalk is long and the corneal extremity 
large, as in E. dovii. 

The sternum and abdomen resemble those of E. dovii^ the anterior 
end of the sternum is depressed, the depression having a convex 
posterior margin, from which a furrow leads back to the abdomen. 
Surface of sternum and abdomen covered with large and distant 
punctae. Abdomen broadly triangular; first segment not dis- 
tinguishable; second, third, and fourth segments each crossed by a 
transverse ridge; third, fourth, and fifth fused, and perhaps also the 
second with them. The sides of the penult segment are less con- 
vergent than in E. dovii. 

This species in all the characters visible in the type-specimen re- 
sembles the genus Euphylax as typified by E. dovii^ excepting in the 

1 Proc. U. S. Nat. Mus., vol. 18, 1898, pis. 12-23. 

« Arch. f. Natarg., vol. 22, pt. 1, 1866, p. 129. 

• Ann. Lye. Nat. Hist. N. Y., vol. 7. 1860, p. 226, pi. 6. fig. 5. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 167 

areolation and ornamentation of the central part of the carapace 
which indicate an aflBnity with GaUinectes. 

EUPUYLAX FORTIS, new tpecic*. 

Plate 64. figs. U-13. 

Type-locaUty. — Banana River, Costa Rica ; tenth f ossilif erous zone 
below tlie uppermost one of the section. Probably equivalent to 
Oatun formation. Miocene series. D. F. MacDonald, collector. 
October, 1911. Station 5882A. Cat. No. 324233, U.S.N.M. 

Measurements. — ^Length of body (approx.), 43.5 mm.; length of 
movable finger, 27.3 mm. ; width of the sternum between the coxae 
of the chelipeds, 24.5 mm. 

Uolotype. — One specimen showing a part of the lower surface 
and the right' cheliped. The abdomen resembles that of an imma- 
ture female or is possibly that of a male. 

Sternum broad, surface rough with coarse punctae and fine reticu- 
lating lines ; anterior part depressed and with a median groove lead- 
ing back to the abdomen much as in E. caUinectias ; the ridge just 
in front of the abdomen is more transverse fhan in that species. 

Abdomen broadly triangular; it is impossible to tell which seg- 
ments, if any, are fused; terminal segment subequilateral ; surface 
of sixth and seventh segments like that of the sternum, of fourth 
and fifth segments covered witli a low, confluent granulation. 

Ischium of external maxilliped with a longitudinal groove, the 
surface on the inner side of the groove more raised than on the 
outer side. 

Cheliped elongate. A cross section of the arm is shown and a 
portion of its lower surface; this last has a broad longitudinal de- 
pression through tlie middle, and the surface near the margins, at 
least, is coarsely granulate. 

The general outline of the fingers can be made out and the sur- 
face of some of the prehensile teeth. The fingers are elongate, as 
in the usual Portunid, and gradually taper, ending in slender black 
tips which cross each other. The prehensile teeth are large, thick, 
dark-colored, and very irregular, the one at the base of the dactylus 
being the largest; they appear to fit close together. The cheliped 
is larger and stronger in proportion to the size of the body than in 
any recent species of Portunid. 

The generic position of this species is problematical ; in the width 
of the sternum it resembles Euphylax; in the strong teeth of the 
digits it approaches ScyUa^ while the groove on the lower side of the 
arm joint is unique. 



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168 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

GATUNHDAE, new family. 
Characters of the type genus, Oatvmd. 

6ATUNIA, new gennB. 

Carapace of the customary Cancrid outline, that is, transversely 
oval, with a narrow, dentate front (between the orbits) ; orbits nar- 
row, with a forward inclination; antero-lateral margins arcuate^ 
longer than the posterolateral, and armed with 8 teeth, including 
the orbital tooth; postero-lateral margins strongly convergent- 
Genital region very narrow. Carapace without transverse ridges. 

Outer maxilliped witli the ischium greatly elongate, and longi- 
tudinally grooved. 

Chelipeds massive; palms thick, not flattened on the inner side^ 
devoid of ridges on the outer side. 

Last pair of feet with the propodus and dactylus flattened and 
broadened to form a swimming organ as -in the Portunids. 

Abdomen of the male with the third, fourth, and fifth segments^ 
fused. 

This genus resembles the family Cancridae in the form of the 
carapace, front and orbits; while the swimming paddles and the ab- 
domen are like those of the Portunidae. The chelipeds approach 
those of the genus Scylla in their massiveness, long fingers and lack 
of costae, but the absence of spines gives them the appearance of 
many of the Xanthidae. 

Type of the genus. — Gatunia proavita Kathbun. 



GATUNIA PROAVFTA, nc 

Plates 54-56; plate 58, figs. 16 and 17. 

Type-locality. — Gatun formation, near Gatun Dam, Panama Canal 
Zone. Miocene series. Collected by one of the workmen and 
shipped by D. F. Macdonald. Station 5659. One specimen (holo- 
type) , nearly complete. Cat. No. 324289, U.S.N.M. 

Measurements. — Length of carapace, from tip of submedian teeth^ 
133.2 mm.; from median sinus, 128.3 mm.; width, between tips of 
teeth of posterior pair, 182.5 mm.; width between teeth of penulti- 
mate pair, the same. 

Holotype. — Carapace about 1^ times as wide as long; antero-lateral 
margin strongly arched, cut into 7 strong teeth, besides the tooth 
at the outer angle of the orbit; teeth similar in shape, having a convex 
posterior and a concave anterior margin, tip acute; the 7 teeth in- 
crease in size from the first to the fifth and then diminish to the 



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OBOLOGY AND PALEONTOLOGY 07 THB OANAL ZONE. 169 

seventh, which is the most spinif orm. The orbit has a strong tooth 
at the onter and at the inner angle and one less prominent between; 
below the inner angle there is a narrow tooth more advanced than 
those above. Frontal region, between the orbits, with a concave 
dorsal surface; anterior border cut into 4 teeth, those of the middle 
pair nearer to eadi other than they are to those of the outer pair, and 
separated by a shallower sinus; teeth of inner pair oblong, with 
rounded end; those of outer pair thicker. Postero-lateral margins 
nearly straight; posterior margin slightly convex and thickened. 

Surface covered with a pavement of fine, flattened granules, and 
with less numerous and unequal punctae visible to the naked eye. 
The depression between the gastric and branchial regions is well 
marked except anteriorly, the hepatic region not being defined. 
Genital region very narrow, longer than wide. Cardiac and intestinal 
regions incompletely outlined. 

Neither the eyes nor the antennae are visible. 

Epistome subtriangular, prolonged downward at the middle in an 
atute angle; thence a small button-hole groove runs obliquely back- 
ward. Palatal ridge strong except anteriorly where it is low and 
blunt. Pterygostomian region granulate, densely so near the buccal 
cavity. On the sternum a furrow runs obliquely forward from the 
coxae of the chelipeds to the median line. 

The abdomen of the male is broad and at the base reaches to the 
coxae of the last pair of feet; there is only one segment visible be- 
tween the carapace and the third segment; it is probably the second, 
while the first is hidden under the carapace much as in the Portunid 
genus CaUinectes; the second is of nearly even length throughout 
its width ; the third, fourth and fifth are fused, but their extent is 
indicated by indentations in the lateral margins and by a short groove 
at the middle; the 3d segment is produced sideways beyond the 
2d and 4th and its margins are very convex ; margins of the 4th to 
6th segments, inclusive, taken together are slightly convergent, those 
of the 4th a little convex ; 6th segment about If times as wide as long; 
terminal segment nearly as long as the preceding, subtriangular, end 
rounded. 

Chelipeds very stout, in general smooth, there being no ridges nor 
spines. The surface is finely granulate and punctate, the granules 
a little higher than on the carapace. The inferior, anterior margin 
is the only margin of the merus visible; it is smoothly rounded. 
Carpus massive, with a broad tooth at inner angle. Chelae unequal, 
thick, broadly rounded above and below without marginal lines; 
right or larger manus about 1^ times as long as high, left or smaller 
manus about 1^ times as long as high ; next the articulation of the 
larger palm with the dactylus there is a large lobe or tooth directed 
toward the end of the dactylus, as in Scylla. The digits each have 2 



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170 BX7LLBTIN 103, UNITED BTATE8 NATIONAL MITSEXTM. 

longitudinal punctate impressions on the outer surface, the larger 
dactylus is more arched than the smaller; the fingers cross each 
other a little before the tips, at least in the smaller chela; the end 
of the larger immovable finger is broken off; prehensile edges armed 
with large irregular, separated teeth, the basal tooth of the larger 
dactylus being of enormous size and directed obliquely backward 
as in ScyUd. The distal half or two-fifths of the fingers is dark- 
colored, also the prehensile teeth. 

The 8 pairs of ambulatory legs are only partially preserved; they 
would, if extended, reach about to the middle of the manus of the 
chelipeds; the first five segments are stout; the propodus, of which 
only impressions exist, has a groove through the middle and appears 
to be flattened; the dactylus can not be made out with any degree of 
certainty; the swimming-feet, or those of the last pair, are very 
broad, tiie carpus is as broad as long; the propodus is about twice 
as long as broad; the dactylus is lanceolate-oval, about 2^ times as 
long as broad. 

Another specimen (paratype a) lacking the dentate border of tBe 
carapace, and all appendages except the coxal joints of the legs, was 
taken from the Gatun formation, Gutun Locks, by D. F. MacDonald, 
May, 1911. Station 6900. Cat. No. 324241, U.S.N.M. 

A much smaller specimen (paratype h) which is identified as 
probably belonging to this species is labeled : " Near Gatun. Mio- 
cene. Rev. G. Rowell. Cat. No. 113706, U.S.N.M." It is probably 
from the Gatun formation. A portion of the left side of the cara- 
pace is preserved, showing the base of the 3 posterior of the antero- 
lateral teeth; on the imder side is shown the margin of the buccal 
cavity, fragments of a maxilliped and the base of the cheliped. A 
separate specimen (paratype c), from the same locality, is the left 
manus, somewhat crushed, proximally incomplete and lacking the 
propodal finger, but with the base of the dactylus attached, including 
the first or large, rounded tooth. 

A fragment of a finger bearing 3 teeth (paratype d) is referred 
here; the smooth outer layer is almost gone except a few bits near 
the teeth; it was taken from the 85-foot cut on north side of big 
swamp on relocated line of the Panama Railroad, 1^ to 2 miles be- 
yond Camp Cotton toward Monte Lirio; Gatun formation; Miocene 
series: D. F. MacDonald and T. W. Vaughan, collectors, 1911; 
Station 6030; Cat. No. 324242, U.S.N JM. 

I refer here with doubt a curved fragment of a thick-shelled species, 
which has a large tooth occupying half its surface. It may belong 
near one of the articulations. It was taken at Station 6033&, in the 
upper part of the lowest bed, Gatun section; Gatun formation; 
Miocene series; MacDonald and Vaughan; 1911; Cat No. 324286, 
U.S.N.M. 



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4 

GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 171 

The larger claw of this species is strikingly like that described by 
A. Milne Edwards^ under the name ScyUa mioheUni from Sceanx, 
near I>ou6, France, in the Miocene sheU deposits of the shell-marl 
of Anjou. M. Milne Edwards f omided the species on the claw alone. 
It is very likely congeneric if not conspecific with the form here 
described. 

Family XANTHIDAE. 

CABPIUUS, ipedM. 

Plate 58, fig. 22. 

Locality. — ^Panama Canid Zone. Foraminiferal marl and coarse 
sandstone about 200 yards south of southern end of switch at Bohio 
Bidge station, relocated line Panama Bailroad. Upper part of 
Culebra formation. Oligocene series. D. F. MacDonald and T. W. 
Vaughan, collectors. 1911. Station 6025. Cat. No. 824243, U.S.N.M. 

Material. — ^Piece of propodal segment of ambulatory leg on left 
side of crab. Length 17 mm., greatest width 7.2 mm., least width 
5.7 mm., proximal thickness 4.5 mm., distal thickness 8.7 mm. Viewed 
dorsally, the anterior margin is slightly convex, the posterior faintly 
concave. Viewed edgeways, the upper surface is longitudinally con- 
vex, and the lower surface concave. Cross section oval. Surface, 
except for accidental breaks, smoothly rounded, without ridges, 
furrows, or tubercles. 

In its smoothness and general form, resembles the propodus of the 
first ambulatory leg of Carpiliw coraJMnus (Herbst*), for which 
reason I venture to attach the name OarpUius to this fragment. 

HBTBRACTAEA LUNATA (MUiie Bdwftids and Laeu). 

Plate 68, figs. 7-9. 

PUutnnus lunatus Mn.NB Edwabds and Lucas, d*0rbigny'8 Toy. Am6r. M6r^ 
VOL 6, 184S, p. 20; vol. 9, aUas, 1847, pi. 9, fig. 2. 

Locality. — Costa Rica : City of Port Limon. Port Limon forma- 
tion. Pliocene series. Dr. L. A. Wailes, collector. Station 4269. 
Cat. No. 824265, U.S.N.M. 

Distribution. — ^Recent, San Diego, California, to Chile. 

Material. — One specimen showing distal portion of outer surface 
of larger palm, with proximal half of dactylus (showing all sur- 
faces) attached. This must have belonged to a small individual 
with carapace about 15 mm. wide. The fossil is crushed and the 
tips of the tubercles are lacking. The shape of the two segments so 

> Hlttolre d€t Cnutacte podophthalnudret foMilet, Paris, 1861, p. 180, pL 8, flgt. 8, 8i.. 
•Natnrg. <L Krabben a. Krebw, toL 1, 1788, p. 188, pL 6, flf. 40. 

8870*— IS— BuU, 108 ^12 



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172 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

far as it is preserved accords with that of recent specimens; the 
dactylus is more deflexed than it could be if the immovable jfinger 
were preserved in sUu. The tubercles of the palm are arranged 
in general as in recent individuals, and slight divergences are attribu- 
table to individual variation. The dactylus has 6 punctated grooves, 
and the prehensile tooth situated at a little distance from the base is 
present; the three uppermost ridges bear some tubercles, the outer 
ridge two tubercles, the upper one three tubercles, and the inner ridge 
one tubercle followed by several crenulations. 

PANOPBUS ANTEPUBPUREUS, IMW tptim. 

Plate 58, figs. S-IL 

Type-locality. — ^Panama Canal Zone. From near Mount Hope in 
ditch through swampy groimd. About one-fourth mile from present 
sea beach, 6 to 8 feet above high tide. Pleistocene series. D. F. 
MacDonald, collector. April, 1911. Station 5850. 

Types.'-CB.t. No. 824246, U.S.N.M. 

Material. — 18 dactyli of stronger chela from right side; 9 dactyli 
of stronger chela from left side ; 4 dactyli of weaker chela from left 
side ; one propodal finger of weaker chela from right side. 

With one exception these digits belonged to small individuals; the 
exception, a ri^t dactylus 12.8 mm. long, is made the holotype. 

The dactyli are very much like the corresponding parts of P. 
pwrpureu8 Lockington,^ a recent species ranging from Ix)wer Cali- 
fornia to Peru. The only differences are as follows : The fingers are 
a little longer, slenderer, and straighter; the large basal tooth of the 
stronger chela is closer to the articulation with the manus; there is no 
coarse granulation on the basal portion of the dactyli, as there is on 
the living form. Most of the specimens are purplish-blue except at 
the tip. 

The propodal finger also is slenderer than in P. purpureus; the 
lower groove of the outer surface is nearer the lower margin. As this 
finger was not attached to a dactylus, one cannot be positive that it 
belongs to the same species as the dactyli. 

PANOPBUS TRroENTATUS, new spcdM. 

Plate 58, figs. 12-15. 

Type-locaUty. — ^Panama Canal Zone. From near Mount Hope in 
ditch through swampy ground. About one-quarter mile from present 
sea beach, 6 to 8 feet above high tide. Pleistocene series. D. F. 
MacDonald, collector. April, 1911. Station 5850. 

Types.— Csit. No. 324244, U.S.N.M, 

iProc. California Acad. Scl., vol. 7. 1876 (1877), p. 101. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 175 

Material. — Eleven dactyls and three propodal digits of the right 
chela, all detached. One of the dactyls is made a holotype. 

Measvrements, — Length of longest dactyl 9.2 mm., height 3 mm, ; 
length of holotype 6.1 mm., height 2.3 mm. 

Holotype. — I have chosen a small specimen for holotype because 
it is the best preserved. The dactyl is rather slender for the major 
chela of a Panopeid, but it has the general character of Paaiopeua 
and its allies. The prehensile edge has 3 enlarged teeth, placed as 
follows, beginning at the proximal end : 1 1., 2 s., 1 1., 3 s., 1 1., 7 s., tip^ 
The proximal tooth is pf the customary backward-pointing type,, 
and is of moderate size compared to the basal tooth of living Pan- 
opeids. The second of the large teeth is of nearly the same size,, 
more pointed, and directed downward ; the third large tooth is define 
itely smaller than the others. The small teeth are unequal and shal- 
low. The longitudinal depression either side of the teeth is punc- 
tate; there are 3 other punctate furrows, one external, one internal 
and the other dorsal but nearer the outer side; just within the proxi- 
mal half of the dorsal furrow there is a marginal line of granules. 

Paratypes, — The number of small teeth intervening between the 
larger teeth of the prehensile edge vary as follows, beginning at the 
proximal end : 1-3, 4-6, 6-8. 

One can not be sure that the propodal digits referred here belong 
to the same species as the dactyli. They too have 3 large teeth, which 
are subequal and are separated by small teeth as follows, beginning 
with the proximal large tt)oth: 2, 4-5, 4-6. Proximal end broken 
oflf in each case, but in one instance a small tooth is visible proximal 
to the first large tooth. When an immovable finger is applied against 
a movable finger of complementary size, the large teeth of the former 
shut into the sinuses distal to the corresponding large teeth of the 
latter. There are 6 longitudinal punctate depressions on each pro- 
podus, one adjacent to the teeth on either side, one external, one in- 
ternal, and two inferior. 

PANOPEUS, species. 

Plate 66, figs. 8 and 9. 

Locality. — From the four feet of dark, stratified tuff and clay^ 
inmiediately overlying the lower limestone bed. Las Cascadas sec- 
tion. Upper part of Culebra formation. Oligocene series. D. F. 
MacDonald and T. W. Vaughan, collectors. 1911. Station 6019ft. 
Cat. No. 324254, U.S.N.M. 

Material. — One dactylus of right cheliped, with proximal end in- 
complete. Dactylus very broad at base in proportion to its length,, 
also unusually thick. Length, 9.2 mm.; width, 4.7 mm.; thickness, 
3 mm. In outer view the upper margin is much curved, the surface 



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174 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

is much worn, but there is a line of punctae visible through the mid- 
dle; there is a large subbasal tooth directed obliquely backward, and 
bounded posteriorly by a deep, wide groove ; the tooth does not pro- 
ject beyond the general lower margin, but it may have done so when 
entire. On the distal portion of the outer edge of the prehensile sur- 
face there are a few shallow crenulations. 

This specimen comes nearer to Panopeua than to any other known 
genus; from P, chiUiisis Milne Edwards and Lucas ^ it differs in 
its greater width, in the large tooth originating higher up on ihc 
outer surface and in the groove behind it being deeper and more 
extensive. 

EURTTIUM CRENULATUM. new ipedes. 

Plate 64, figs. 8 and 9. 

Type-locality. — Panama Canal Zone. From near Mount Hope in 
ditch through swampy ground. About one-quarter mile from pres- 
ent sea beach, 6 to 8 feet above high tide. Pleistocene series. D. F. 
MacDonald, collector. April, 1911. Station 5850. 

Holotype.— Cat No. 324253, U.S.N.M. Dactylus of right chela, 7.6 
nam. long. This has the general shape of a Euryfium finger, but the 
large basal tooth of the prehensile edge is inserted higher up, its 
oblique base running posteriorly well up on the outer surface; the 
tooth is directed strongly backward and is broadly roimded at the 
extremity. It is followed by about ten low teeth, the first and third 
of which are the larger. Somewhat abc^ve the middle of the inner 
jund of the outer surface there is a longitudinal depression containing 
:a row of a few punctae. There is a deep groove on the upper sur- 
face and just within it but higher up there is a row of separated 
granules or crenulations; they point outward, that is, a side view of 
them may be obtained by looldng down on the top of the finger. 
The ridge just outside the dorsal groove is proximally microscopically 
granulate. 

This finger can not be referred to E. affine^ or E, tinstani^ of the 
Panamian fauna on account of the elevation and direction of the 
basal tooth, and the row of gianules on the upper edge. 

Family GONEPLACIDAE. 

Subfamily Prionoplacinae. 

EURTPLAX CULEBRENSIS. new species. 
Plate 66, figs. 13 and 14. 
Type-locality, — Panama Canal Zone. Top part of limy sandstone 
below upper conglomerate, near foot of stairs, Gaillard Cut. Upper 

1 D'Orblgny's Voy. Am6r. M«r., vol. 6, pt. 1, 1843, p. 16 ; vol. 9, atlas, 1847, pi. 8. flg. 2. 
'Panopeus affinia Streets and Kingsley, Bull. Essex Inst., voL 9, 1877, p. 106. 
•Rathbun, Proc. Biol. Soc. Washington, vol. 19, 1906, p. 100. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 175 

part of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6012^?. 

H olotype,— C2it. No. 324226, U.S.N.M. Propodus of right and 
major cheliped with only the base of the immovable finger remaining. 
Palm swollen. In side view the upper and lower margins are arcu- 
ate, the palm narrowing considerably toward the wrist. Surface 
smooth. At the distal end above the lower margin there is a deep 
groove which is prolonged on the finger. The palmar tooth which 
overlaps the dactylus is present; also 4 prehensile teeth of the fixed 
finger, arranged in 2 transverse rows on the upper surface, the outer 
tooth of the distal pair being much the largest. 

Measurements. — ^Length of palm, to sinus, 6.6 mm.; height, 4.3 
nmi. ; thickness, 2.7 mm. 

This specimen has the general form of E. nitida Stimpson,^ a Re- 
cent species occurring on the coast of the Gulf of Mexico and the 
West Indies. I have no example of the Panamian species, E. polita 
Smith* for comparison. E. nitida is considerably larger than the 
fossil form, the fixed finger is somewhat wider at the base but it has 
the 4 basal teeth similarly disposed ; the proximal end of the upper 
margin is thinner and more acute than in E. culebrensis. 

Subfamily Hexapodinae. 

Living representatives of this subfamily are restricted to the Indo- 
Pacific region. 

Genus THAUMASTOPLAX Mienu 

Thnumastoplax Miers, Ann. Mag. Nat Hist, sor. 5, vol. 8, 1881, p. 261. 
The generic position of the species placed here has to be deter- 
mined by the characters discernible in a dorsal view. As in Thau- 
mastopldXj the shape of the carapace is subrectangular with the 
antero-lateral comers roimded off; the second ambulatory leg is 
stronger than the first and third. Of the other Hexapodinae, or 
Goneplacids with only 3 pairs of walking legs, Hexapus de Haan* 
is more subcylindrical and has the three legs of subequal size; Larnb- 
{lophallus AJcock* has smaller orbits; Hexapldx Doflein' has very 
oblique orbits seen from above, while Paeduma Kathbun* (=i4m<?r- 
phopns Bell ^) is said to be almost cylindrical. 

* Ann. Lye. Nat. Hist. New York. vol. 7, 1859, p. 60. 

* Trans. Connecticut Acad. Scl., vol. 2, 1870, p. 168. 
' Fauna Japon., 1833, p. 5 ; 1885, p. 35. 
«Journ. Asiat. Soc. Bengal, vol. 69, 1900, p. 829. 

•'Wiss. Ergeb. deutschen Tlefsee-Exped. VtA^via, 1898-99, vol. 6, 1904, p. 122. 
•Proc. Biol. Soc Washington, vol. 11, 1897. p. 168. 
^ Jour. Linn. Soc London, Zool., vol. 8, 1858, p. 27. 



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176 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

THAUMASTOPLAX PRIMA, imw ipectof. 

Plate 66, figs. 15-18. 

Type-locality. — ^Panama Canal Zone. Las Cascadas section, Gail- 
lard Cut. From lowest fossiliferous bed; third bed below lowest 
limestone beds separated by rows of nodules. Lower part of upper 
half of Culebra formation. Oligocene series. D. F. MacDonald 
and T. W. Vaughan, collectors. 1911. Station 6020a. 

Measurements. — ^Length of carapace, paratype, 12.2 mm. (ap- 
prox.) ; width, 18 mm. (approx.) The holotype is a little wider, 
about 19.6 mm., but the length can not be measured as the front part 
of the carapace is not visible. 

Holotype and paratype. — Carapace about 1^ times as broad as long, 
longitudinally very convex, from side to side nearly horizontal. Up- 
per surface not bordered by a definite line; H-shaped depression in 
the center of the carapace deep; surface deeply punctate, punctae 
crowded near the margins, sparse near the middle. Fronto-orbital 
distance about two-fifths as great as the extreme width of the cara- 
pace. Antero-lateral margins long, arcuate ; postero-lateral margins 
subparallel; posterior margin slightly convex. Front deflexed and 
widening a little from the base of the eyestalks downward. The 
orbit is about as wide as the narrowest part of the front and is filled 
by the eyestalk; its upper margin is transverse. 

Chelipeds short, when flexed scarcely reaching beyond the outer 
end of the orbit; carpus very large, convex, smooth, and punctate; 
chela small, not much longer than carpus and considerably nar- 
rower; finger shorter than palm; the end of the finger is, however, 
not visible. The merus of the first leg reaches quite to the end of 
the carpus of the cheliped, its upper margin has a row of small 
conical tubercles or granules, and there is a cluster of granules near 
the articulation with the carpus. The merus of the second leg 
reaches a little beyond that of the first, and is very much stronger; 
it also has a superior row of granules and a few granules on the 
outer surface; carpus elongate, about half as long as merus; pro- 
podus as wide as the greatest width of the carpus; only a portion 
of it is visible. Third leg very much shorter and slenderer, its 
carpus reaching little beyond the merus of the second pair ; its merus, 
as well as that of the second pair, is longitudinally grooved. 

The above description is made from two specimens from the same 
place. Each specimen was enclosed in a nodule which is broken in 
two. The holotype shows the upper surface of the carapace (ex- 
cept the front part) , portions of the left cheliped, and of the 3 legs 
of both sides. The nodule is not large enough to have included the 
whole of the legs in their extended position. Cat. No. 324227, 
U.S.N.M. The paratype shows the carapace only; there is no trace 



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GBOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 177 

of chelipeds or legs; the nodule is very little wider than the carapace. 
Cat, No. 824228, U.S.N.M. 

Family GECARCINIDAE. 

CARDISOMA GUANHUm LatMllto. 

Plate 64, figs. 2-4. 

Cardisoma Quanhumi Latbett.t.e, Encyc. M§tli., Hist Nat., Insectes, vol. IG, 
1825, p. 685. 

Locality. — Costa Rica : City of Port Limon. Port Limon forma- 
tion. Pliocene series. Dr. L. A. Wailes, collector. Station* 4269. 
Cat No. 324263, U.S.N.M. 

Material. — ^Left propodal finger of cheliped, with extremities lack- 
ing; length 18.5 mm. Pieces of the outer crust remain along the pre- 
hensile teeth, around the distal end, and on the lower part of the 
proximal end, where it shows the characteristic scaly granulation of 
the species. 

Distribution of Recent Material. — From Bahamas and Florida 
Keys to Brazil; Bermudas. 

Family OCYPODIDAE. 

UGA MACRODACTTLtJS (Milne Edwudi md Lqcu). 

Plate 64, fig. 7. 

Oelasifnus macrodactylus Miunc Edwabds and Lucas, d^Orbigny's Yoy. 
Am^. M^r., vol. 6, 1843, p. 27 ; vol. 9, atlas, 1847, pi. 11, fig, 8. 

Locality. — ^Panama Canal Zone. From near Mount Hope in ditch 
through swampy ground. About one-quarter mile from present sea 
beach, 6 to 8 feet above high tide. Pleistocene series. D. F. Mac« 
Donald, collector. April, 1911. Station 5860. Cat. No. 824251, 
U.S.N.M. 

Material. — ^A single dactylus, 6.7 mm. long, of an ambulatory leg 
corresponds with that of a recent specimen from Costa Rica. The 
dactylus is rather slender, regularly tapering and strongly curved 
on its concave as well* as on its convex margin, it has 6 longitudinal 
grooves separated by as many smooth rounded ridges; toward the 
homy tip these ridges are themselves guttered by a narrow groove. 

Distribution of Recent Material. — From Guaymas, Mexico, to Val- 
paraiso, Chile. 

BRACHYRHTNCHA, family, geniu, and species indeterminable. 

Plate 64, fig. 6. 

Locality. — ^Panama Canal Zone. Las Cascadas section, Gaillard 
Cut, From lowest fossiliferous bed. Third bed below lowest lime- 



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178 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

stone beds separated by rows of nodules. Lower part of upper half 
of Culebra formation. Oligocene series. D. F. MacDonald and 
T. W. Vaughan, collectors. 1911. Station 6020a. Cat. No. 324294, 
U.S.N.M. 

Material. — A specimen 9.3 mm. long and 7.7 mm. wide, which re- 
sembles the proximal part of the right manus of a crab. The outer and 
lower surfaces are exposed, but the outer layer of shell has almost dis- 
appeared. The surface gradually ascends to a line a little below the 
middle where a blunt longitudinal ridge is formed. Just below the 
upper margin, and also just within the lower margin on the inner 
side, tjiere is a narrow furrow. 

The blunt crest is suggestive of the Portunidae. 

BRACHTRHTNCHA, family, genus, and spedes indeterminable. 

Plate 64. fig. 5. 

Locality. — Panama Canal Zone. Las Cascadas section, Gaillard 
Cut From lowest fossiliferous bed. Third bed below lowest lime- 
stone beds separated by rows of nodules. Lower part of upper half 
of Culebra formation. Oligocene series. D. F. MacDonald and 
T. W. Vaughan, collectors. 1911. Station 6020a. Cat. No. 324258. 
U.S.N.M. 

Material. — Two specimens, each embedded in a nodule, of a frag- 
ment which appears to be the merus segment of an ambulatory ]fig 
of a crab. The surface is flat, the lateral margins are arcuate and 
there is a shallow longitudinal depression near one edge. The shape 
is very suggestive of the Portunidae and yet they do not closely 
resemble any known species. 

Superfamily OXYRHYNCHA. 
Family PARTHENOPIDAE. 

PARTHENOPE PANABfENSIS, new spedet. 
Plate 66, figs. 10 and 11. 

Type-locality. — ^Panama Canal Zone. Las Cascadas section. 
From fifth or topmost limestone. Emperador limestone. Oligocene 
series. D. F. MacDonald and T. W. Vaughan, collectors. 1911. 
Station 6019^^. Cat. No. 324257, U.S.N.M. 

Measurements. — ^Length of arm measured along lower margin (in- 
complete), 24.6 mm.; distal width, measured on inner, lower surface, 
6.6 mm. ; minimum width, on the same surface, 4.7 nnn. 

Holotype. — One specimen represented by only the merus joint of 
the left cheliped. This segment is thick; a cross section is quadri- 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 179 

lateral and nearly as broad as long; the proximal end is broken oflF^ 
the distal end is embedded in a hard matrix. The two upper sur- 
faces are each not much more than half as wide as either of the two 
lower surfaces. The upper, inner and outer margins are armed with 
irregular spines and tubercles, the lower margin with tubercles only, 
which are not in a single row ; the surfaces also have a few scattered 
spines and tubercles. The tips of most of the spines are broken off ; 
the largest spines are one on the upper margin near the proximal end 
(of the specimen), and one just above the outer margin in the middle 
third. The distal extremity of the segment widens in a manner sug- 
gestive of the propodus of a Parthenopid, but it is not triangular- 
prismatic as customary in the propodal segments. 

PARTHENOPE PLEI8T0CENICA. new ipedes. 

Plate 61, figs. 10 and 11. 

Type-locality. — Panama Canal Zone. From near Mount Hope in 
ditch through swampy ground. About one-quarter mile from pres- 
ent sea beach, 6 to 8 feet above high tide. Pleistocene series. D. F. 
MacDonald, collector. April, 1911. Station 5850. Cat. No. 324247, 
U.S.N.M. 

Holotype.-^'PvoipoAvil finger of right and major chela ; lower proxi- 
mal portion missing. Lower margin, so far as it remains, nearly 
straight, tip upturned; upper margin broad at the proximal end 
and armed with 5 imequal blunt teeth; the first or that nearest the 
palm is broadly united with the second which is much larger >. third 
of similar size and shape to the second and well separated from it; 
fourth much smaller and fifth minute. Four lateral rows of punctae, 
one either side of the prehensile teeth, and one through the middle 
of the inner and of the outer surface. The posterior half of both 
surfaces is sparsely covered with prominent and very unequal 
granules. 

Parthenope excavata (Stimpson)^ of which there is an example 
from Panama in the United States National Museum collection has 
a similar prehensile surface, but the lateral face is shorter and higher 
and is granulate all over outside. 

EXPLANATION OF PLATP^S. 

Plate 54. 

Oatunia proavita Ruthbun, holotype, dorsiil view, X g. 

Plate 55. 

Oatunia proavita, holotype, ventral view, X \. 

^Lamhrut excavatus Stimpson, Ann. Lye. Nat. Hist. New York, vol. 10, 1871, p. 08. 



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180 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Plate 56. 

Gatunia proavita^ holotype. 

Pig. 1. Anterior view, X |. 
2. Posterior view, X |. 

Plate 57. 

Fig. 1. Natantla, indetermiuable, lateral view of pleon. X 2. 

2. Axius reiiculatus Rathbun, holotype, outer view of propo<lus of left 

chellped, X 3. 

3. Impression of same. 

4. Macrobrachium, specie.s, Cat. No. 324256, outer view of propodus of left 

cheliped, X 3. 

5. Upper view of same. 

C. LcucosUia hananensis Rathbun, paratype a, arm joint, X 3. 

7. TA'ucoHilia hananams, paratype &, arm Joint, X 3. 

8. LcuconiUa hanancnftiif, holotype, arm Joint, X 3. 

0. Macrohravhiumf, species, Cat. No. 324248, segment of large chelli>ed, 
X 34. 
10. Axinst, spofies, Cat. No. 324250, scaphocerite, X 3^. 

11. Goniochcle'i armaia Rathbun, holotyi)0, outer view of dactylus of left 

(jheliped, X 2. 

12. Lower view of same, X ::. 

13. NtphropH costaivs Ratlibun, holotype, dactylus of left chellped. Inner 

view, X 31. 

14. Dorsal view of same, X 3^. 

15. Nephrops contatuSf paratype a, dactylus of left chellped, dorsal view, 

X 3i. 

16. Js'ephrops costatus, paratype c, propodal finger of left chellped, dorsal 

view, X 3i. 

17. Nephrops coslatus, paratype 6, dactylus of left chellped, dorsal view, 

X 3i. 

18. VeirolisihcH aviius Rathbun, holotype, palm of left chellped, dorsal view, 

X 3i 

19. Inner view of same, X 3i. 

20. Ventral view of same, X 3i. 

■21. VavhycheJcs lalits Rathbun, holotype, propodus of left chellped, ventral 

view. X 3i 
"22. Dorsal view of same. 

23. Pachychclcs latns, paratype, left chela, dorsal view, X 3i. 

24. Calappa costaricana Rathbun, holotype portion of propodus of left 

chela, outer view, X 3. 

25. Nephrops, species, Cat. No. 324240, dactylus of right chellped, prehensile 

edge, X 3i. 

26. Dorsal view of same, X 3^. 

27. Mursilia ecristata Rathbun, holotype, right chellped, outer view, X 3. 

Plate 58. 

Tig. 1. Calappella quadrispina Rathbun, holotype, impression of carapace, X 8. 

2. Carapace of figure 1, X 8. 

3. Pwnopeus antepurpureus, Rathbun, holotype, dactylus of right chellped, 

X 8i. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 181 

Fio. 4, 5. Panopeu9 antepurpureus, paratypes, dactyl! of right chellpeds, X 3^. 
6. Panopeus antepurpureus, paratype. Immovable finger of right cheliped, 
X 3i 
7-11. Panopeus antepurpureus, paratypes, dactyli of left chellpeds, X 8i. 

12. Panopeus tridentatus Rathbun, paratype, dactylus of right cheliped, X 31. 

13. Panopeus tridentatus, paratype, propodal finger of right cheliped, X 3). 

14. Panopeus tridentatus, holotype, dactylus of right chelipetl, outer view, 

X Sh 

15. Upper edge of same, X Si. 

16. Fragment of Oatunia proavita ft. Cat No. 324286, showing lobe in profile, 

X 2. 

17. Flat surface of same, X 2. 

18. Callinnassa hilH Rathbun, paratjrpe 6, sixth segment of pleon, X 2. 

19. Callianassa hilU, holotype, merus and carpus of right cheliped, X 2. 

20. Callianassa JiUH, paratype a, right cheliped, X 2. 

21. Mursia macdonnldi Rathbun, holotype, left palm, X 2. 

22. CarpUius, specios, Cat. No. 324243, piece of propodus of a left ambulatory 

leg, X 2. 

Plate 50. 

Fig. 1. Callianassa oralis Rathbun, holotype, left cheliped, X 2. 

2. Callianassa oralis, paratype a, propodus of left cheliped, X 2. 

3. Impression of figure 1, X 2. 

4. Callianassa ovalis, paratype 6, carpus of left cheliped, X 2. 

5. Callianassa, species, Cat. No. 324277, manus of left cheliped, X 3i. 

6. Callianassa lacunosa Rathbun, holotype, left manus, distal view, X 3. 

7. Outer view of same, X 3. 

8. Inner view of same, X 3. 

9. Callianassa Uwunosa, paratype, left manus, inner view, X 3. 
10. Outer view of same, X 3. 



11. Distal view of same, X 3. 



Plats 60. 



Fio. 1. Callianassa nioinensis Rathbun, holotype, propodus of right cheliped, 
lower view, X 3. 

2. Upper view of same, X 3. 

3. Outer view of same, X 8. 

4. Callianassa elongata Rathbun, holotype, propodus of left cheliped, upper 

view, X 2. 

5. Outer view of same, X 2. 

6. Lower view of same, X 2. 

7. Leucosiidae?, indeterminable. Cat No. 324136, dactylus of left chela, 

outer Aiew, X 3^. 

8. Upper view of same, X 8}. 

9. Callianassa scotti Brown and Pilsbry, holotype, left manus, lower view, 

X li. 

10. Callianassa scotti, paratype, Oat No. 2259, left propodus, upper view, 

X 11 

11. Outer view of same, X IJ. 

12. Callianassa scotti. Cat. No. 824279, left manus, distal view, X li. 
18. CaUianassa tenuis Rathbun, holotype, left manus, inner view, X 8. 
14. Outer view of same, X 8. 



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182 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Plate 61. 

Fio. 1. CalHanassa crassa Uiithlmii, holotype, dactyliis of left chelli)ecl, upper 
view, X 2. 

2. Outer view of same, X 2. 

3. CalHanassa crassoy paratype, dactylus of 'eft ehellped, outer view, X 3. 

4. Calappa flammea, dactylus of right chelipeti, inner view, X 2. 

5. Outer view of same, X 2. 

6. Callianassa spinulosa Rathbun, paratype iu Mus. Acad. Nat Sci. Phila., 

propodus of left ehellped, outer view, X 2. 

7. Callmnassa spinulosa^ holotype and impression, left propodus, outer 

view, X 2. 

8. Distal view of same holotype, X 2. 

9. Inner view of same, X 2. 

10. Parthenope pleistocenica Kathbun, holotype, propodal finger of right 

chela, upper view, X 3i. 

11. Outer view of same, X 3i. 

12. CaUianassa stridens Rathbun, holotype, left manus, outer view, X S. 

13. Upper view of same, X 3. 

14. Inner view of same, X 3. 

15. Callianassa crassimana Rathbun, holotype, propotlus of. left cheliped, 

inner view, X 2. 

16. Impression of same, and piece of finger, X 2. 

17. Holotype laid against impression of same, outer view, X 2. 

18. Mursia obscura Rathbun, holotype, carapace, X 3. 

Plate 62. 

Fig. 1. Callianassa magna Rathbun, holotyi>e. dactylus of right chelii)ed, Inner 
view, X li. 

2. Outer view of same, X 1^. 

3. Upper view of same, X 1^. 

4. Callianassa quad rata Rathbun?, carpus of left cheliped, upper view. X 2. 

5. Ijovver view of same, X 2. 

6. Outer view of same, X 2. 

7. CalHaiwssa quadrata, holotype, left manus, distal view, X 2. 

8. Inner view of same, X 2. 

9. Outer view of same, X 2. 

10. Callianassa quadrata^ paratype a, left manus, outer view, X 2. 

11. Inner view of same, X 2. 

12. Distal view, X 2. 

13. Callianassa quadrata, paratype e, right manus, outer view, X 2. 

14. Callianassa quadrata, paratype d, left manus, outer view, X 2. 

Plate 63. 

Fio. 1. Callianassa ahbrcviata Rathbun, holotype, right manus, inner view, X 3. 

2. Outer view of same, X 3. 

3. Distal view of same. X 3. 

4. Impression of same, X 3. 

5. CaVinnasm ahbrcviata, paratype t, piece of prop<Klus of left cheliped, 

outer view, X ,*i. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 183 

Fig. 6. CaUianassa abbreviata, paratype a, left manus, outer view, X 3. 

7. Heteractaea lunata (Milne Edwards and Lucas), right chela, Cat. No. 

824266, outer view, X 3i. 

8. Upper view of same, X 31. 

9. Heteractaea lunata, right chela of Recent specimen, Cat. No. 2140, X 3i. 

10. CallianaMa vaugTiani Rathbun, paratype b, portion of propodus of loft 

cheliped, outer view, X 2. 

11. CaUianassa vaughani, paratype a, left manus, outer view, X 2. 

12. Inner view of same, X 2. 

13. CaUianassa vaughani, holotype, right chela, outer view, X 2. 

Plate 64. 

Fio. 1. Arenaens, species, Cat. No. 324252, left movable finger, outer view, 
X 3*. 

2. Cardisoma guanhumi I^atreille, propodal finger of left cheliped, outer 

view, X 3. 

3. Lower view of same, X 3. 

4. Upper view of same, X 3. 

5. Brachprhyncha, indeterminable, Cat. No. 324258, merus of ambulatory 

leg, X 3. 

6. Brachyrhyncha, indeterminable, Cat. No. 324294, right manus, X 3. 

7. Uca macrodactylus (Milne Edwards and Lucas), Cat. No. 324251, dacty- 

lus of ambulatory leg, X 3i. 
10. CalUanassat, species. Cat. No. 324280, merus -of right cheliped (?), 
view, X 31. 

9. Upper view of same, X 31. 

10. CaUianassa r, siiecies. Cat. No. 324280, merus of right cheliped (?), 

X 31. 

11. Euphyla^ fortis Rathbun, holotype, right side, showing cross sections of 

merus of cheliped and of two legs, X 11. 

12. Front view of same, showing remains of fingers, X 11. 

13. Ventral view of same, X 14 

Plate 65. 

PiQ. 1. CaUincctcHy species. Cat. No. 324255, manus and carpus of left cr.elliieil, 
outer view, X 11. 

2. Upper view of same, X 11. 

3. Euphylax caUinectias Rathbun, holotype, dorsal view, X 11. 

4. Posterior view of same, X 11. 

5. Ventral view of same, X 11. 

6. Anterior view of same, X 11. 

7. CaUincctes, species. Cat. No. 324268, distal third, tip missing, of immov- 

able finger, X 3. 

Plate 66. 

Fig. 1. Callinectes declivis Rathbun, holotype. propodus of left cheliped, upper 
view, X 11. 

2. Inner view of same, X 11. 

3. Outer view of same, X 11. 

4. Hepatus oJUliensis Milne Edwards, Cat. No. 324235, dactylus of right 

chela, inner view, X 31. 

5. CaUincrtes rctiatlatns Rathbun, holotyi)e, propodus of right chelipe<l, 
• upper view, X 11. 



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184 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Fio.6. Inner view of same, X 1|. 

7. Outer view of same, X IJ. 

8. Panopeus, species, Cat No. 324254, dactylua of riglit cheliped, upper 

view, X 3. 

9. Outer view of same, X 8. 

10. Parthenope panamensis Rathbun, liolotype, merus of left chelipecU outer 

view, X 11. 

11. Inner view of same, X IJ. 

12. Hepatu8, species, Oat No. 824239, dactylus of left cheliped, outer view, 

X 2. 

13. Euryplax culebrenais Rathbun, holotype, propodus of right chelipeO, 

upper view, X 8i. 

14. Outer view of same, X 81. 

15. TTiaumastoplax prima Rathbun, paratype, carapace, X 2. 

16. Impression of same, X 2. 

17. Thauniastoplax prima, holotype, X 2. 

18. Impression of same, X 2. 



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INDEX. 



Pace. 

aW>reTiata. CaUlanassa 125, 120, 

187, 147, 182, 183 
acanthurus panamenBe, Macrobra- 

chium 131 

affine, Eurytium 174 

afDnis, Panopeus 174 

AroorpbopuB 175 

an^^ata, Goniochelc 155 

Anomura 134 

antepurpureus, Panopeus 128, 130, 

172, 180, 181 

Arenaeus mexicanus 165 

species 128, 130. 1G5, 183 

armata, Gonlochele? 125, 120, 154, 180 

Mursla 150 

Porcellana 135 

armatus, Petrollstbes 135 

AsUcura 132 

avltns, PetroUsthee 127,120.134,180 

Aziidae 135 

Axlus 123, 136 

retlculatas 125,120,185,180 

species 128, 129, 136, 180 

baharaensis, Cancer 153 

Petrochlrus 153 

Balanus concavus rariseptntus 126, 129 

eburneus 127, lu28, 129 

glyptopoma 127, 129 

( Hesperibalanus ?) , s p e - 

cles 124, 125, 129 

bananensis, Leucosilia ._ 127, 130, 161, 180 

bocourtl, Calllnectes 166 

boayierl, Petrochinis 129, 153 

Brachygnatba 162 

Bracbyrbyncha 162 

family, genus and 
species indetermin- 
able 125, 130, 

177, 178, 183 

Bracbyura 154 

Calappa 160 

costaricana 127, 130, 156, 180 

flammea 128, 130, 157, 182 

gallus 156 

rurcheri— 124, 130. 157 

Calappella 130, 157 

quadrisplna 125, 130, 157, 180 

Calappidae 155 

callforniensls, Petrochlrus 153 

Callianassa 124, 186 

abbrevlata 125, 120, 137, 

147, 182, 183 



Pai^. 

CalliasaBsa crassa_.12e, 120, 137, 152, 182 
crassimana 125, 120, 

137, 141, 182 
elongata 125, 120, 186, 

180, 146, 181 

hilli 126, 120, 187, 148, 181 

lacunosa 124, 120, 

136, 138, 181 
magna— 125, 120, 187, 151, 182 
moinensis 126, 120, 

186, 142, 181 
ovalls — 125, 120, 186, 187, 181 

pellucida 151 

quadrata 124, 125, 120, 

137, 145, 182 
scotti.. 124, 125, 120, 136, 140, 

142, 148, 145, 146, 181 

species 120, 152, 181 

species 125, 120, 153, 183 

spinulosa 124, 125, 120, 

186. 148, 182 

stlmpsoni 148 

strldens 125, 120, 

187, 151, 182 
tenuis.- 126, 120, 186, 144, 181 

toulal 124, 120, 187, 146 

vaughani 125, 126, 120, 

137, 148, 183 

Callianassidae 130 

Calllnectes 163, 166, 169 

bocourtl 106 

decllvls -- 127, 130, 

162, 164. 183 

exasperatus 166 

reticulatus 125, 130, 

163, 164, 183 

sapldus 163 

species 1^6, 130, 164. 183 

tozotes ' 165 

callinectias, Bupbylax 127, 130, 

166, 167, 183 

Oancer (Astacus) jamalcensls 131 

bahamensls 153 

flammeus 157 

gallus 156 

Cancrldae 123 

Cardlsoma 123 

guanhumi 127. 130. 177, 183 

Carpillus 171 

corallinus 171 

species 125, 180, 177, 181 



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INDEX. 



chilensis, Panopeus 174 

chlUensis, Hepatus 128,129,155,183 

concavuft rariseptatus, Balanus — 126, 129 

corallinus, Carpilius 171 

coBtaricana, Calappa 127, 130, 156, 180 

costatufi, Nephrops- 128, 120, 132, 133, 180 
crassa, CalliaDassa . 126, 129, 137, 152, 182 
crasslmana, CalllanasBa 125, 

129. 137. 141. 182 
crenulatum, Eurytium ._ 128, 130, 174, 183 

culebrensis, Earyplax 125, 130, 174, 184 

Decapoda 131 

decliyis, CaUinectes. 127, 130, 162, 164, 183 

dovii, Eaphylax 166 

Dromiacea 154 

Dromiidae 154 

eburneuB, Balanus 127, 128, 129 

ecristata, MursUla 126, 130, 160, 180 

elongata, Callianassa 125, 

129, 136, 139, 146, 181 

Euctenota mezicaDa 165 

Euphylax 123, 166, 167 

calUnectias 127, 

130. 165. 167. 183 

dovll 166 

fortls 127, 130, 167, 183 

Euryplax 123 

culebrensis 125, 180, 174, 184 

nltlda 175 

pollta 175 

Eurytium 123, 174 

afflno 174 

crenulatum 128, 180, 174, 183 

tristani 174 

exasperatos, CaUinectes 166 

oxcavata, Parthenope 179 

cxcavatus, Lambrus 179 

flammea, Calappa 128, 130, 157, 182 

flammeus. Cancer 157 

fortls. Euphylax 127,130,167,183 

Galatholdea 134 

fl^alathlna, Porcellana 135 

galathinus, Petrolisthes 135 

gallus, Calappa 156 

Cancer 156 

Gatunia . 130, 168 

proavita 126, 

130, 168, 179, 180, 181 

Gatunlldae 108 

(Jf*carcinidno 177 

Gelasimus macrodactylus 177 

glyptoporoa, Balanus 127, 129 

Goneplacldae 123, 174 

Goniochele 154 

angulata 155 

armata 125. 120, 154, 180 

granulatUH, Petrocbirus 153 

grosslmanuB, Pachychelcs 134 

guanhuml, Cardisoma— _ 127, 130, 177, 183 

Hepatus 123 

chiliensls 128, 129. 155, 183 

species 125, 155, 184 

Hesperlbalanus, species 124, 125, 129 



Heteractaea 123 

lunaU 127. 180. 171, 18» 

Hexaplax 175 

Ilexapodlnae 123, 175 

Hexapus 175 

hllli, Callianassa 126, 129, 137, 148, 181 

Homaridae 182 

injudicata, Lepas 126, 129 

jamaicense, Macrobrachium 181 

jamaicensls, Cancer (Astacus) 181 

Jurlnel, LeucosUia 128,180,161 

lacunosa, Callianassa 124 

129. 136. 188. 181 

Lambdophallus 176 

Lambrus excavatus 179 

latus, Pachycheles 127,129.184,180 

Lepas injudicata 126, 129 

Leucosiidae 161 

genus and species inde- 
terminable 128, 

130. 162. 181 

Ix'ucosilia 128 

bananensis ... 127. 180. 161^ 180 

Jurinei 128.130.161 

lunata, Heteractaea 127, 180, 171,188 

lunatus, Pilumnus 171 

macdonaldi, Muraia 126.130,158,181 

Macrobrachium 181 

acanthurus p a n a - 

mense 181 

jamaicense 131 

mexicanumu 131 

species— 126,129,130.180 
vpedes. 128, 129. 182, 180 

macrodactylus, Gelasimus 177 

Uca 128,180.177.183 

mapia, Callianassa. 125, 129. 187, 151, 182 

Malacostraca 131 

mexicana. Euctenota 166 

mexlcanum, Macrobracblum 181 

mesicanus. Arenaeus 165 

Palaemon 181 

mlchelinl. Scylla 171 

molnensis, Callianassa 126. 129, 

186. 142. 181 

Mursla 128. 160 

armata 159 

macdonaldi 126, 180. 158. 181 

obscura 124. 180,159, 182 

Mursllla 160 

ecristata 126, 180, 160. 180 

Natantla 181 

family, genus, and species 

indeterminable 129, 181, 180 

Nopbrops 188 

costatus 128, 129, 132. 180 

species 128,129,133.180 

Xeptunus ^^ 163 

iiltlda. Euryplax 175 

obsoura. Mursla 124,180.159,182 

Ocypodidae 177 

ovalls, Callianassa 125.129, 

180, 187. 181 
Oxyrhyncha 178 



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Ill 



Page. 

Oxystomata 155 

I^cbycbeles 128 

grossimanus 184 

latus 127, 129, 184, 180 

Paediima 175 

Pagurldae — 158 

PalaemoD mexlcanus 181 

Palaemonidae .^ 181 

I>a]iaiDe]i0e, Macrobracblam acan- 

thurus 181 

I>aiiameD8i8, Parthenope. 126, 180, 178, 1^ 

Panopeus . 178,174 

affinis 174 

antepurpureus 128, 180, 

172. 180. 181 

chllenaiB 174 

purpureua 172 

spedea 125, 180, 173,184 

trldcntatus 128, 180, 172, 181 

Parthenope ezcayata 179 

panamenalB- 126, 180, 178, 184 

pleistocenlca 128, 180. 

179, 182 

Parthenopldae 178 

Pecten 124, 141 

pellndda, Callianasta 151 

Persephona 162 

Petrochlrus 124 

bahamensls 158 

bonvieri 129, 153 

callfomlensiB 158 

granalatos 158 

Petrolislbes 123 

armatua 185 

avltus 127, 129. 184^180 

galatblntis 186 

Pllumnus lunatus 171 

plelstoceDiea, Parthenope 128, 

180. 179. 182 

polita, Buryplax 175 

Porcellana armata 135 

galatbina 135 



Fagf.. 

Porcellanidae 134 

Portunidae 128, 162 

Portunus 163 

sanguinolentus 103 

prima, Thaumastoplax _. 125, 130, 175, 184 

Prlouoplaclnac 174 

proavlta, Gatunia 126, 

130. 168. 179. 180. 181 

purpureas, Panopeus 172 

quadrata, Calllanassa 124 

125. 129. 187. 145. 182 
Quadrisplna, Calappella.. 125, 180, 157, 180 
rarlseptatus, Balanus concavus— 126. 129 

Reptantla 132 

retlculatus, Axius 125, 129, 135, 180 

Callinectes 126, 

130, 103, 164, 183 

sanguinolentus, Portunus 163 

sapidua, Callinectes 163 

Bcotti, CalUanaasa 124, 125, 129, 

136, 140, 142, 148, 145, 146, 181 

Scylla 167, 168, 169, 170 

mlcbellni 171 

vpinulosa, CalUanasaa 124, 

125, 129, 136, 143, 182 

stimpsoni, Calllanassa 148 

stridens, Calllanassa 125, 

129, 137, 151, 182 
tenuis, CalUanasaa. 126,129.136,144,181 

Thalasslnidea 185 

Thaumastoplax 175 

prima — 125.180,175,184 

toulal, Calllanassa 124, 129, 137, 146 

toxotea, Callinectes 165 

trldentatus, Panopeus — 128, 130, 172, 181 

tristani, Burytium 174 

Uca macrodactylus 128, 130, 177, 183 

vaughanl. Calllanassa 125, 

126,129.137.148,183 

Xanthidae 171 

lurcheri, Calappa 124,130,157 



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SMITHSONIAN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 
Bulletin 103 f^ ' - 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE GANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



CIRRIPEDIA FROM THE PANAMA 
CANAL ZONE 



By HENRY A. PILSBRY 
Oj the Academy of Natural Sciences, Philadelphia 



Eitraci from Bulletin 103, page* 185-188, with Plate 67 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1918 



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SMITHSONIAN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 
Bulletin 103 



CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



CIRRIPEDIA FROM THE PANAMA 
CANAL ZONE 



By HENRY A. PILSBRY 
Oj the Academy of Natural Sciences, Philadelphia 



Extract from Bulletin 103, pages 185-188, with Plate 67 




WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1918 



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CIEKIPEDIA FKOM THE PANAMA CANAL ZONE. 



By Henry A. Pilsbry, 
0/ the Academy of Natural Scieficcs, Philadelphia, 



The small number of barnacles collected show the following rela- 
tions. The Pleistocene and Pliocene species are identical with recent 
Atlantic coast and Caloosahatchie Pliocene species, respectively, and 
are distinctively Atlantic forma Of the Oligocene and Miocene 
species, one has relatives in both oceans, another only in the Pacific; 
the affinities of the third being doubtful. 

BALANUS EBURNEUS Govld. 

Balanus ebumeus Ph^sbby, BuU. U. S. Nat. Mus., No. 93, 1916, p. 80, pi. 24, 
figs, l-lo, 2, text-figs. 14 and 15, and synonymy. 

This recent species is represented by four individuals from Station 
5867, the wall being preserved complete, but without opercular plates. 
The smooth surface and the closely, regularly septate parietal tubes 
are characteristic, the septa forming cells about equal in length and 
breadth, throughout the tubes. 

Locality^ and geologic occurrence. — ^They are from a dark mud 
formation about 10 feet above the present sea level, near lower end 
of Gatun Locks. Pleistocene series. D. F. MacDonald, collector. 
April, 1911. Cat. No. 324297, U.S.N.M. Five specimens, from Sta- 
tion 5868, from Mount Hope, in swamp ditch, in black mud forma- 
tion; 'Pleistocene; D. F. MacDonald* 1911. Cat. No. 324290, U.S.N.M. 

Ten specimens of the same were taken at Station 6038, also from 
black mud f romlower end of Gatun Locks. Pleistocene series. D. F. 
MacDonald, collector, 1911. Cat. No. 324293, U.S.N.M. 

BALANUS 6LTPTOPOMA Pflsbrr. 

Plate 67. figs. 1-3. 

Balanus oonoavui glyptopoma Pilsbby, Bull. U. S. Nat. Mus., No. d3, p. 102, 
pL 21, fig. 2 ; pi. 22, figs. 2-2c. 

The walls only of several groups growing on oysters and scallops 
were collected. They agree with the above species described from 
the Pliocene of the Caloosahatchie Biver, and show some additional 
characters, notably the color. The radii are broad. The parietes are 
weakly ribbed longitudinally, the intervals in the best preserved in- 
dividuals being of a deep livid brown color, the low ribs white. The 
parietal tubes are crossed by many septa, down to the base; these are 
a little less regular than in B. ehvmeus^ most of the cells being longer 
8870/— 18 185 



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186 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

than wide, exactly as in the type of B. glyptopomcu There are 16 
tubes in the rostrum of one of the specimens figured (fig. 2). The 
basis is profusely porous, the pores septate (fiig. 3, left hand and 
middle individuals of group shown in fig. 1). The middle specimen 
of this group is 31 mm. high, the rostro-carnial diameter about 
22 mm. 

B. glyptopoma was described as a subspecies of B. concavus Bronn, 
but it differs from that by the closely septate parietal tubes, and is 
evidently a distinct species. In B. concaws the tubes are filled up 
near the summits, and are open, with very few septa below, or none 
in the American subspecies. The Miocene form formerly referred to 
B. glyptopoma is a distinct subspecies of B. conoavw. 

Location and geologic occurrence. — ^The specimens are from Sta- 
tion 5903, across Chagres River and about 200 to 225 feet above it, 
top of hill opposite Alhahuela, in a gray tuf aceous limestone, Cat. 
No. 324298, U.S.N.M. and Station 5906a, 50 to 75 feet below 5905, 
Cat. No. 324299, U.S.N.M. Both collected by D. F. MacDonald. 
Upon mentioning to Dr. William H. Dall that I had identified a 
Pliocene barnacle from these Stations, he kindly informed me that 
'^ both are above the Oligocene strata and separated from the latter 
by an unconformity. They are doubtless Pliocene. 5906a is the 
lower of the two horizons.*' 

Mexico. From the Sayula District of Chiapas, on the Arroyo 
Chapapoapam. Pliocene series. Dr. C. W. Hayes and others, col- 
lectors ; 1911. Station 5886. One specimen, without opercular valves. 
Cat. No. 324291, U.S.N.M. 

BALANUS CONCAVUS BARISEPTATUS. new ratopMlM. 

Plate 67, fig. 4. 

In form this barnacle is somewhat cylindric with contracted sum- 
mit in the adult stage, convexly conic when young. The orifice is 
ovate. The walls are only slightly roughened longitudinally. The 
carinolateral compartments are narrow, the parietes about one-third 
as wide as the lateral compartments. The radii are wide with 
oblique summits, without pores; the articulating edges being cr^iu- 
lated. The parietal tubes have very few, irregularly scattered, trans- 
verse septa. There are 29 tubes in the rostrum of the type-specimen. 
Another, of equal size, has 17 tubes in the lateral, 6 in the carino- 
lateral compartment. 

Length, 27 mm. ; carino-rostral diameter, 22.5 mm. ; lateral diame- 
ter, 21 mm. In the largest individual exposed the rostrum is 85 mm. 
long. 

This form is represented by a group of about 16 individuals grow- 
ing upon and largely concealing a single old one of about 37 tool 
basal diameter. Probably three generations are present. They were 



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QBOLOOY AND PALEOKTOLOaY OF THE OAKAL ZONE. 187 

in a tufaceous limestone. The tubes of the walls are solidly filled 
with calcite. 

This was at first thought to be a smooth form of Balanua concavtts 
Bronn, but on cutting it the parietal tubes were found to be much 
more nimierous. In a specimen of B. concavvs from the British Bed 
Crag (Pliocene), No. 12058, U.S.N.M., there are 19 tubes in the 
rostrum, which is 65 mm. long. Most of these tubes have transverse 
septa at long, irregular intervals, but in some places near the edges 
the septa are rather close, though irregular. 

The relation of this form to B. concavua can not be exactly esti- 
mated until the opercular plates are found. It may -be an ancestral 
form of concavus or a distinct species. Meantime, it is readily recog- 
nizable by the characters of the compartments. 

Locality and geologic occurrence. — ^Panama Canal Zone. From 
85 foot cut, just on north side of big swamp, on relocated line Panama 
Bailroad, li to 2 miles beyond Camp Cotton toward Monte Lirio. 
Oatun formation. Miocene series. D. F: MacDonald and T. W. 
Vaughan collectors, 1911. 1 cluster; Cat. No. 324292, U.S.N.M. 

BALANUS (HBSPBRIBALANU87). tpedM. 

A small, conic barnacle having a basal diameter of about 7 or 8 
mm. is represented by several compartments and one incomplete 
specimen, without opercular valves. The walls are smooth except for 
slight ripples parallel to the base. They are solid, having no parietal 
tubes. The compartments are rather thick for so small a barnacle, 
and when parted the articulating edges of the radii and the opposed 
sutural surfaces are seen to be conspicuously crenulated. The basis 
is calcareous, thin, and seems to have radial threads on its inner 
face. 

These characters indicate a species of the subgenus HesperibaUmusy 
or possibly ScUdohalanvs. Neither group has been recognized 
hitherto in American tertiary deposits, or in the recent faunas of the 
Panamic region or western Atlantic. The specimens do not seem 
characteristic enough to serve as the basis of a new species, though 
they can not, I think, be referred to any described form. 

Locality and geologic occurrence. — ^They were collected by Mac- 
Donald and Vaughan in the " lowest f ossilif erous bed, the third below 
the lowest limestone bed. Las Cascadas section, Gaillard Cut. Lower 
part of upper half of Culebra formation. Oligocene.'' Station 6020a, 
Cat. No. 824296, U.S.N.M. 

A single valve was taken one- fourth mile south of Empire Bridge, 
from lower dark clay beneath lower conglomerate, lower part of 
Culebm formation, Oligocene; Station 6012a; Cat No. 824296, 
U.S.N.M. 

8870'--18— Ball. 103 13 



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188 BULLETIN 103, UNtTSD BTATB8 NATIONAL MUSEUM. 

LBPA8 INJUDIGATA. b«w ipmIm. 
Plate 67, fig. 5. 

This species is based upon a calcareous plate believed to be the 
scutum of a Lepadid barnacle. It is thin, trapezoidal in outline, the 
basal border straight, almost equal to the length, and a little con- 
tracted or narrowly bent in. The tergal extremity is broad and 
somewhat convex. The occludent margin is almost straight. The 
surface has the curvature of an ordinary Lepas^ such as L. anatif^rny 
and is sculptured with coarse, unequal concentric folds, with, to- 
wards the growing edges, some rather fine concentric striation. 

The fossil is imperfect at the tergal end, but if restored according 
to the lines of growth it would be about 26 mm. long; width 23 mm. 

That the fossil has been correctly interpreted is by no means cer- 
tain. If Lepadid, as believed, the very obtuse tergal end probably 
indicates a small, transversely placed tergum, not running between 
scutum and carina, or perhaps none. Either condition would denote 
greater specialization than the modem genus Lepaa. However this 
may be, the fossil is specifically recognizable by its form and sculp- 
ture, and we must await the finding of further material to reveal 
its nature. 

LocalUy and geologic occurrence. — ^The holotype is No. 324448, 
n.S.N.M. It was found by MacDonald and Vaughan in a section of 
the bluffs exposed along the Panama Railroad. Relocation, about 
3,500 feet south of Gatun Railroad Station, in bed No. 6033&, Gatun 
formation.. Miocene series. 

EXPLANATION OF PLATE 67. 

Fio. 1. BaUmua glyptopoma PUsbry. Lateral view of group from Station 5903. 
2. Rostral view of a spedmen of B. glyptopona growing on Peoten, Station 
8. Basal view of fig. 1. 

0903, the outer lamina of the wall removecL Length of roBtnmt 
16 "i"^ 
4. Balanus concavus rariiepiatus Pilsbry. Type. 
5. Lepas injudioata Pilsbry. Type. 



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U. 8. NATIONAL MUSEUM 



BULLETIN 103 PL. 67 




Panama Cirripedes. 

For explanation of plate sec pace iss- 



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SMITHSONIAN INSTITUTION I lil j ^ 

UNITED STATES NATIONAL MUSEUM » WV • / ^ 
Bulletin 103 

CONTRIBUTIONS TO THE GEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



FOSSIL CORALS FROM CENTRAL 
AMERICA, CUBA, AND PORTO RICO, 
W^ITH AN ACCOUNT OF THE AMER- 
ICAN TERTIARY. PLEISTOCENE, AND 
RECENT CORAL REEFS 



By THOMAS WAYLAND VAUGHAN 

Custodian of Madreporaria, United States National Museum, and Geologiit in charge 
of Coasud Plain Investigations, United States Geological Survey 



Extract from Bulletin 103, pages 189-524, with Plates 68-152 






WASHINGTON 

GOVERNMENT PRINTING OFFICE 

1919 



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SMITHSONIAN INSTITUTION 

UNITED STATES NATIONAL MUSEUM 

Bulletin 103 



CXDNTRIBUTIONS TO THE CEOLOGY AND PALEON- 
TOLOGY OF THE CANAL ZONE, PANAMA, AND 
GEOLOGICALLY RELATED AREAS IN CEN- 
TRAL AMERICA AND THE WEST INDIES 



FOSSIL CORALS FROM CENTRAL 
AMERICA. CUBA, AND PORTO RICO. 
WITH AN ACCOUNT OF THE AMER- 
ICAN TERTIARY. PLEISTOCENE, AND 
RECENT CORAL REEFS 



By THOMAS WAYLAND VAUGHAN 

Cuuoiian tf hiajnptraria, Uniud Smut National iluuum, and GttUgitt in tkargt 
tf CtmuJ Plain Invtsiigmions, Uniud Stout Gioltgical Saro$y 



Extract from Bulletin 103, page* 189-524, with Plate* 68-152 




WASHINGTON 

GOVERNMENT PRINTING OFHCE 

1919 



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CONTENTS. 



Introduction 189 

Geologic correlation by means of foesil corals 190 

Geologic history of the upper Eocene and latw coral faunas of Central America, 

the West Indies, and the eastern United States 193 

Eocene 193 

Brito formation, Nicaragua ! . . . 193 

St. Bartholomew limestone 193 

Jackson formation and Ocala limestone 195 

Concluding remarks on the Eocene 198 

Oligocene 198 

Lower Oligocene 198 

Middle Oligocene 199 

Antigua formation 199 

Pepino formation of Porto Rico 203 

Limestone above conglomerate near Guantanamo, Cuba 204 

Basal part of Chattahoochee formation in Georgia 205 

"Coral limestone" of Salt Mountain, Alabama 206 

San Rafael formation of eastern Mexico 206 

Tonoei, Panama 207 

Serro Colorado, Arube 207 

Concluding remarks on the middle Oligocene 207 

Upper Oligocene 208 

Culebra formation 208 

Emperador limestone 208 

Anguilla formation 209 

Cuban localities 210 

Tampa formation of Florida ' 211 

Conduding remarks.on the upper Oligocene 211 

Miocene • ; 212 

Bowden marl. 212 

Santo Domingo 213 

Cuba 218 

Baracoa and Matantas 218 

La Cruz marl • 218 

Florida 219 

Alum Bluff formation 219 

Middle and South Atlantic States 220 

Costa Rica 221 

Panama '. 221 

Colombia 221 

Concluding remarks on the Miocene 221 

Pliocene 222 

Caloosahatchee marl, Florida 222 

Limon, Costa Rica 223 

Carriso Creek, California 223 

PleiBtocene 225 

m 



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IV COISTTENTS. 

Geologic history of the upper Eocene, etc.— Continued. Pa«©. 
Summary of the stratigraphic and geographic distribution of the Tertiary 

and Pleistocene coral-faunas of Central America and the West Indies. . . 226 

Table of stratigraphic and geographic distribution of species 228 

Conditions under which the West Indian, Central American, and Floridian 

coral reefs have formed, and their bearing on theories of coral-reef formation . 238 

Definition of the term* 'coral reef" 238 

Ecology of reef-forming corals 240 

Hypotheses of the formation of coral reefs 241 

Tests of coral-reef hypotheses 246 

Criteria for recognizing shift in position of strand line 246 

Criteria for measuring the amount of vertical shift in strand line, and 
for determining the relative ages of terraces and the physiographic 

stage attained by a shore line 247 

Criteria for ascertaining the rdle of corals as constructional agents 248 

Solubility of calcium carbonate in sea water 260 

Effects of wind-induced and other currents in shaping coral reeia 251 

Criteria for determining the effect of glaciation and degladation on 

the development of living reefs 252 

Amount of vertical displacement of strand line by glaciation and 

deglaciation 252 

Rate of growth of corals and length of post-Gladal time 253 

Effect of loweriug of marine temperature on reef onals during 

glaciation '. 254 

Valley-in-valley arrangement and cliffed i^urs 256 

American Tertiary and Pleistocene reef corals and coral reefe 258 

Eocene reef corals of St. Bartholomew 259 

West Indian middle Oligocene reels 259 

Antigua 259 

Porto Rico 260 

Cuba 261 

West Indian and Panamanian upper Oligocene ree^ 262 

Anguilla 262 

Canal Zone 262 

West Indian Miocene reef corals 263 

West Indian Pleistocene reefe 263 

Tertiary and Pleistocene reef corals and coral reefs of the United States. 265 

Southeastern United States 265 

Plicene reef corals from Carrizo Creek, California 271 

Living coral reefs of the West Indies, Florida, and Central America 271 

. Antigua-Barbuda Bank 273 

St. Martin Plateau 275 

St. Croix Island 278 

ViiginBank 279 

Cuba 280 

Bahamas : 291 

Bermudas 293 

Flotida 297 

CampecheBank 298 

Honduran reefs... 300 

Mosquito Bank 300 

Some other West Indian Islands 301 

Brazil and Aigentina 301 

Atlantic coast of the United States north of Florida 303 

Tyi>es of West Indian and Central American littoral and suUittoral 

profiles and their relations to coral reefe 303 



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OONTBNTS. V 

Conditions under which the West Indian, Central American, and Floridian 

coral reefo have formed — Continued. Page. 
Living c<»al reels of the West Indies, Florida, and Central America— Contd. 
Submeiged banks north of the coral reef zone in the western Atlantic 

Ocean 305 

Summary of the conditions under which the American f ossO and living 

reefa formed 305 

Coral reefs of the Pacific Ocean 306 

Great Barrier Reef of Australia 306 

New Caledonia ^. 308 

Fiji Islands 309 

Society Islands 311 

Tahiti 311 

Smaller islands of the Society group.- 312 

Atolls 313 

Conclusions 319 

Bearing of these conclusions on hypotheses of the formation of coral reefs 325 

Suggestions as to future investigations...!....^ 329 

Systematic account of the faunas 333 

Class Anthozoa 333 

Madreporaria Imperiorata 333 

Family Seriatoporidae .' 333 

Genus Stylophora '. 333 

Podllopora 342 

Ifadrads ' 345 

Family Astrocoenidae 345 

Genus Astrocoenia 345 

Stylocoenia 351 

Family Ociilinidae 352 

Genus Oculina 352 

Archohelia 352 

Family Eusmiliidae 354 

Genus Asterosmilia 354 

Stephanocoenia 356 

Didiocoenia 360 

Eusmilia 361 

Family Astrangiidae 361 

Genus Cladocora 361 

Family Orbicellidae 362 

Genus Orbicella 362 

Solenastrea ^ 395 

Antiguastrea 401 

Stylangia 410 

Sepastrea 411 

Family Faviidae 412 

Genus Favia 412 

Favites 414 

Croniastrea 416 

Maeandra 417 

Leptoria 421 

Mftnir>inft 421 

Thysanus 423 

Family Mussidae 424 

Genus Syzygophyllia 424 



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VI OONTBNTS. 

Systematic account of the {aunas--Contiiiued. 

ClasB Anthozoa — Continued. race. 

Madreporaria Fungida * 425 

Family Agarididae 425 

Genus Trochoseris 425 

Genus Agarida. 426 

Pavona 430 

Leptoseris 431 

Pironaatraea 432 

Siderastrea 436 

Family Oulastreidae 458 

Genus Gyathomorpha 454 

Diploastrea 469 

Madreporaria Perforata 479 

Family Eupsammiidae 479 

Genus Balanophyllia 479 

Family Acroporidae 479 

Genus Acropora 479 

Astreopora 483 

Actinacis 486 

Family Poritidae 488 

Genus Goniopora 488 

Pontes 498 

Class Hydrozoa 507 

Order Ilydrocoralliftae 507 

Family Milleporidae 507 

Genus Millepora 507 

Explanation of plates 507 



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FOSSIL CX)RAI5 PROM CENTRAL AMERICA, CUBA, AND 
PORTO RICO, WITH AN ACCOUNT OP THE AMERICAN 
TERTIARY, PLEISTOCENE, AND RECENT CORAL REEPS. 



By Thomas Watland Vauohan, 

Cuitodian qf Madnporaria, UniUd StaU$ NoHorud MuBeum^ and Otologiit in ehatgt 
of CooikU Plain Inv€$tigiUums, United StaUiOeologiealStavey. 



INTRODUCTION. 

The object of the present memoir is to contribute information 
that may aid in deciphering the geologic history of the perimeters 
of the Gulf of Mexico and the Caribbean Sea. Therefore, problems 
of correlation, the physical conditions under which the different 
formations were deposited, and the distribution of land and sea 
during the successive geologic epochs have been particularly in mind. 

The material on which this paper is based is extensive. It 
includes collections made in Panama by Dr. D. P. MacDonald and 
me, working jointly, and by Doctor MacDonald while alone; and Dr. 
Ralph Arnold obtained a small but valuable lot of specimens at 
Empire in the Canal Zone. The collections from Cuba were made 
by Dr. Arthur C. Spencer, Mr. O. £. Meinzer, and myself; the one 
from Porto Rico was made by Mr. R. T. Hill, who also obtained a 
small but valuable lot of specimens in Antigua; the principal col- 
lections from Antigua and Anguilla are the results of my individual 
efforts, and I obtained considerable material in St. Bartholomew, 
but not so much as Qeve got in 1869. There are numbers of small 
lots, as follows: One from Nicaragua, obtained by Dr. C. W. Hayes; 
one from Colombia/ collected by Mr. O. C. Matson; specimens from 
limon, Costa Rica, procured by DoctK)r Wailes and Mr. H. Pittier; 
and specimens from eastern Mexico, obtained by Mr. E. T. Dumble. 
All of the collections mentioned are the property of the United 
States National Museum, having been made in connection with 
official work of some kind, or the material, if privately collected, 
has been presented to the Museiun. Messrs. Matson, WfJles, Pittier, 
and Dumble have presented specimens. My own collecting in 
Antigua, St. Bartholomew, and Anguilla was made possible by a 
minor grant from the Carnegie Institution of Washington, and as a 
result I brought some thousands of specimens to Washington. 

189 



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190 BULLETIN 103, UNITED STATES NATIONAL MX7BBUM. 

These were presented to the United States National Museum by 
the Carnegie Institution, 

Besides haying studied the material indicated, I have twice been 
able to examine all of Dimcan's types preserved in the Museum of the 
Gedogioal Society of London and in the British Museum (Natural 
History), and I heartily thank the (rfScers of those in8tituti<ms for 
the privileges accorded me. In 1904 Prof. A. O. EU^gbom and 
Prof. C. Wiman most generously permitted the Qeve collection 
from St. Bartholomew and Anguilla to be sent to me in Washington. 
This collection contained all of Duncan's types from St. Bar- 
tholomew; and I thank Messrs. Hdgbom and Wiman for the excellent 
opportunity they gave me. Some duplicates from the deve col- 
lection, identified by direct comparison with Duncan's types, were 
procured for the United States National Museum by exchange. 

Opportunities to study the Gabb collection from Santo Domingo, 
divided between the Philadelphia Academy of Natural Sdences and 
the Museum of Comparative Zoology, and the specimen obtained by 
Miss Carlotta J. Maury in Santo Domingo, have been very valuable. 
In fact, as a result of Miss Maury's careful stratigraphic studies in 
that Republic, the stratigraphic relations of the Santo Domingan 
faimas became known. Except retaining a few duplicates, she has 
generously presented to the United States National Museum the 
material obtained by her. 

I wish to thank my associates in the United States Natt<mal Museum 
and in the United States Geological Survey for their helpfulness 
during the prosecution of this study. Mr. W. O. Hazard, of the 
Siu*vey photographic laboratory, made most of the photographs used 
for illustrations, and Miss Frances Wiesser retouched some of them. 

There is almost no literature on the Tertiary fossil corals of 
Central America, Cuba, or Porto Rico. I listed a few Pleistocene 
species obtained by Mr. R. T. TTill at a place IJ miles west of Port 
Ldmon, Costa Rica; * and Felix has recorded from Colombia ' three 
species, as follows: 

OrUcetta (heresiana Felix, probably a synonym of Solenastrea 
houmoni M. Edward and Haime. 

Isastraea turbinata Duncan. 

Stephanocoenia cf. S.fairhanJcsi Vaughan. 

None of these records is further considered in the present paper. 

Toula has described Oculina gatunensis from Gatun (see footnote, 
page 352 of this paper). 

GEOLOGIC CORRELATION BY MEANS OF FOSSIL CORALS. 
That vegetative variation in corafe is great and that without large 
suites of specimens the limits of variation can not be aaoertaLned are 

> Mus. Comp. Zool. Bull., vol. 28, p. 275, 1898. 

s FsUx, J., Ueber elnige fossile Korollen aus Columblen, K. Bayer. Akad. Wi88.,inatb.-phys. Kl.. 
Slteuneiber., vol. 86, pp. 85-03, 1006. 



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OEOUOQY AND PALXOKTOLOOY OF THE OAKAL ZONE. 



191 



two facts so well known to students of Madreporaria that they need 
only to be mentioned. I can not be sure that all of the supposed 
species recorded in this paper as. valid are really vaUd; and perhaps 
in identifying specimens from one locality with species from other 
localities I may not always have discriminated closely enough. I 
am discussing close resemblances and minute differences, for these 
are the basis of correlation within such regional limits as the borders 
of the Gulf of Mexico and the Carribbean Sea, and the recognition 
and proper evaluation of this kind of resemblances and differences 
affect the reliability of the deductions as to age equivalence. I have 
been as careful as I well could be, but I should not like to insist that 
I am always right in these very refined matters of observation and 
of inferences based on such refined observation. In order to mini- 
mize error inherent in such work, I have tried not to rely on one 
species, but on groups of species — ^for instance the species of Orbir- 
ceUa and of Goniopora in both the Emperador limestone and the 
Anguilla formation — and I have utilized the testimony of other 
groups of organisms. 

Comparisons of f aimas according to the percentages of species in 
common may be very misleading. Faunas now living only a short 
distance from each other may have nothing or almost nothing in 
common. In order to illustrate this I am introducing a table of the 
corals obtained in the Cocos-Eeeling Islands by Dr. F. Wood Jones.^ 
Although the list has been published elsewhere, it is not very long 
and strikingly illustrates f aimal phenomena that are of great geologic 
importance. 

LiH of cordU obtained by Dr. Wood Jones in Cocoi-KuUng I$kmd9 and their habitat, 
br.rsbnui6hing; frag.==fragfle; insv.s=ma8sive; pL=^]ate; Iiicn28t.=si]icn2sting. 



Name of speoies and growth-form. 



Habitat. 



Lagoon. 



Barrier pools 

and barrier 

flat. 



Exposed 
barrier. 



5ertefoporaaiMrttZataRlmi2lnger,deIloatelv branched. 

FodUopora buwota Ehrenberg» br., form depends on environment.... 

daiufconito (Esper), br.. rather strong 

verrucosa (Ellis and Solander). stout br 

elegant Dana, strong br., aborted on snrf. 

^ i eyaotfxi M . £:d wards and il ai mo, br., rather strong 

woodjonesi Vaughan, br., rather strong 

Orbicella vrrtipora (Lamarck), msv 



X 
X 
X 



X 
X 



Cjfphastrea mterophthalma (Lamarck), msv 
etlosa fEspor), thin folia... 



&kl7iopoTa lametl.^^ ^-.,, 

LeptaatTca pUTpurea (l)ana), msv 

botiae (M. Edwards and Halmc), msv 

immeTsa Klunzinger, msv 

Favta ttelligera (Dana), msv 

Bpeciosa (Dana), msv. (dead specimen) 

FavUeaabdita (Ellis and Solander), msv 

melicrrum (F:hrenbcrg), msv. (dead specimen) 

Zeptoria phrygia (Ellis and Solander), msv 

Bydnophora mkrocorws (i.amarck), msv. (dead specimen).. 
exua (Fallas), lobato. 



X 
X 
X 
X 



X 

*x" 



* Vaqghan, T. W., Some shoal-water oorala from Kurray Island (AnstnUia), Cocoa-Keeling Iilaiida, 
and Fanning Island, Carnegie Inst. Washington, Pub. 213, pp. 70-72, 1918. 



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192 



BULLETIN 103, XmiTBD 8IATB8 KATIOKAL MU8BX71C 



Li$t ofwrals obtaifud by Dr. Wood Jone$ in Cocot-Kedmg Islandi and their kabiiat — 

Continued. 



Name of spedet and groiTth-form. 



Habitat. 



Lagoon. 



Barrier poob 

and barrier 

Sat. 



Exposed 

barrier. 



FfMQiafufigUtt ^Cinnaeos). free disk, 
teutaria Lamarck, nee disk . 



HerpetoUtka cnuta Dana, free coral.. 

Puvona iamai (U. Edwards and Haime), strong folia. 
fRoMiMfwit (Gardiner), msy 



X 
X 



X 
X 



sariaiM VerrilL msT 

Psammoeofs kaimiana M. Edwards and Haime, msv. 
sp., inorust 



Utfoonedge > 
oTbarrier. 



Sand 



X 
X 



DendropkfUia wUUvi (Oardlner), msv 

dktphtna Dana, liicnjst, base, protob. ooralUtes. 

Aitreopon mfriophthalma (Lamarck), msv 

Montipora Uvit Qiielch, br . , 



krtuo9a (Dana), frag., br. 
ramoia Bernard, fng., br. 

eoeo$en$i» Vaughan, br... 



tpiimota (Lafkiarck), msv. 
sp., lobate columns 



X 
X 
X 



Espedall: 
inner 



X 
X 



Mormit Bernard, msv.. pi. on lower edges... 

/Wto«i( Pallas), thin folia 

iieropora puiekra(BTook), tng.. br 

pkanomt (M. Edwards and Haime)f br 

forma araMca (M. Edwards and Halme) . . 
eoTfmbom (Lamarck), oonrmbose 



X 
X 
X 



X 
X 



spic(^era(Dana),oor7mboae .' 

tiehmeriana (Brueggemann). msv. base, stout br. 
oeelMa (KJunxinger), mBv.iob 



Lagoon 
inlet. 



variahiiit (Klnnzinger ), br 
paUfera (Lamarck), strong br 



PorUtt tottda (Forskftl), msv 

tomaUintit Qravier, msv. 

lieken Dana,incrust 

fUffrefeem Dana, br 

3HtUpora diehoUma Forskal, br. 



ptefvpftfU 
8p.,mcrust . 



f I/a Ehvenberg, strong folia 

rust 

Total number of species according to locality. 



nner margin 
of barrier. 



30 



X 
X 



X 
X 
X 
X 



X 

X 



16 



Of the 23 species found in the lagoon, 3 also occur on the exposed 
barrier, and one of these is so modified to meet surf conditions that 
ordinarily the specimens from the two localities would not be rec- 
ognized as belonging to the same species. Thirteen per cent of the 
lagoon species occur on the exposed barrier; while 18 per cent of 
the exposed-barrier species occur in the lagoon. .These are the 
relations within perhaps half a mile. There are^ 20 species in the 
barrier pools and on the barrier flat. Of these 6 occur within the 
lagoon and 2 were obtained on the exposed barrier; or there are 
30 per cent in common with the lagoon and 10 per cent in common 
with the exposed barrier. When such relations as these prevail 
among the Uving corals of a small group of small islands, what are 
the chances that we should among fossil corals get a large percentage 
of common species? 

The collection listed shows that certain species do occur in all 
three habitats, and, by searching, spots may be found where the 



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GEOLOGY AND PiXBONTOLOGT OF THB OAKAL 202^. 193 

faunas of the different habitats mingle. Corals of the same habitat 
should be compared, or groups of species of the same genera, as I 
have done for Empire (Canal Zone) and Anguilla, where the habitats 
are nearly enough aUke for the same genus to thrive in both. Un- 
less it caQ be established that the habitats are ecologically very 
nearly the same the percentages can not be used safely. 

GEOLOGIC HISTORY OP THE UPPER EOCENE AND LATER CORAL 
FAUNAS OF CENTRAL AMERICA, THE WEST INDIES, AND THE 
EASTERN UNITED STATES. 

EOCBKB. 
BRITO FORMATION, NICARAOUA.i 

Dr. C. W. Hayes collected on or near the Pacific coast of Nicaragua 
the following species: 
Aetrocoenia d^ashiardii Duncan. 
SyzygophyUia hayesi Vaughan. 

ST. BARTHOLOMEW LIMESTONE.* 

I am introducing the name St. Bartholomew limestone for the 
upper Eocene limestones of St. Bartholomew. Description of the 
rock, its stratigraphic relations, and summaries of its faunal char- 
acters are given in the papers referred to in the footnotes. Only 
two species of corals found in the St. Bartholomew limestone are 
actually described in the present memoir, namely: 

Agtrocoenia d'achiardii Dimcan. 

incrustans (Duncan) Vaughan. 

The fossil corals from the St. Bartholomew limestone have been 
specially considered by Dimcan * and myself. Prof. A. O. Hdgbom, 
of the University of Upsala, kindly lent me in 1904 the entire CJeve 
collection from St. Bartholomew, and in 1914 I spent eight days 
studying and collecting on the island. I am combining both the 
Qeve and my collections in the following list, and am adding 
the names of the Jamaican Eocene species, several of which also 

1 For ao acooont of the Brlto formation, see Hayes, C. W., Physiography and geology of region adjacent 
to the Nicaragua Canal route, Oeol. Boo. Amer. Boll., vol. 10, pp. 285-348,1910. Description of the Brito 
formation, pp. 309-313. 

* For aoooants of the geology of St. Bartholomew, see as follows: Cleve, P. T., On the geology of the 
northeastern West India Islands, K. svonska Vet.-Akad. Handl., vol. 0, No. 12, pp. 24-27, 1872. Vaughan, 
T. W., study of the stratigraphic geology * * * of the smaller West Indian Islands, Carnegie Inst. 
Washington Yearbook No. 13, pp. 358-300, 1915; also Yearbook No. 14, pp. 368-373, 1916; [Present status 
of geologic correlation of the Tertiary and Cretaceous formations of the Antilles], Washington Acad. Set. 
Jour., vol. 5, p. 488, 1915; Reef-coral fauna of Carrizo Creek, Imperial County, California, and its signifl* 
outce, U. 8. Oeol. Survey Prof. Pap. 98-T, pp. 362, 363, 1917. 

I Duncan, P. M., On the older Tertiary fonnations of the West-Indian Islands, Oool. Boo. London Quart, 
loom., vol. 29, pp 54»-665, pis. 19-22, 1873. 

Vauf^han, T. W., Soma Cretaceous and Eocene corals from Jamaica, Mus. Comp. Zool. Bull., vol. 84, 
t^. 227-250, 2S5-2S6, pis. 36-41, 1889; A critical review of the literature on the simple genera of the Madre- 
poraria Fungida, with a tentaUve classification, U. S. Nat. Mus. Proc., vol. 28, pp! 371-324, 1905; Study of 
tU stratigraphic geology * * * of the smaller West Indian Islands, Carnegie Inst. Washington Year- 
book No. 13, pp. 358-360, 1915; The reef-coral fauna of Carriso Creek, Imperial County, California, etc., 
V. a Oeol. Survey Prof. Pap. 96-T. pp. 862-363, 1917. 



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194 BULLBTOr 108, UIOXED SIAIBS KAHOKAL MU8BX71C. 

occur in St. Bartholomew. Duncan described Eocene species from 
Jamaica in the papers referred to in the footnotes below.^ 

Eocene eoraUfrom St. Bartholomew and Jamaica. 





tholo- 
mew. 


Jamaica. 




Revised name. 


Cata- 
dopa 
f6rma- 
tion. 


Rich- 
mood 
forma- 
tion. 


Cam- 
brid^ 
forma- 
tion. 


Notes. 


Pheotroekut deoei (Duncan) 


X 

X 
X 

X 








Twbino$€ri9 devetDvaoMiL 


AtttTOtniui vourMtti Duncan 










newspedes 










IVockotifiitta new spedei 








can, iM^Brogniart. 


VW "^ftUFhan, 


X 








£Kytop*ora cofnpreMa Concan 

eofUorta (Leymerle) (fide 
Duncan) 
AHrocoenia duerdeni (Vaughan) 


X 










X 






X 
X 

X 
X 
X 
X 




' V 




incrutUifu (Diincan) 




aupkanocoenia ineruttaiu Don* 


d'achiardU Duncan 






can. 


AntUJia (f) compresw (Duncan) 


( 






(f) ctoW (Duncan) 


1 




drcoi^fma devei DuBCUL ^ 












i 




** Eocene of Jamaica. " 


JMa new speciefll.^ 


X 








new species 2 


X 
X 
X 


.............. 






Goniaatrea variaMU* Vtaieaii 








Maeandra new species 1 






2fanieina areolaU Duncan, ntd 


new species 2.. XX * . x .. 


X 


J 





T ilnnaoDi 
nolReon. 




X 


1 






X 
X 

X 






INpltrfo eo^/)v<ieetls Vaag^L 


eoi^erticosta var. columnarit 
(Vaugban) 
TroekoaeriM ealadwpen^iit Vaughan 














ma/of (Duncan) 

mvuf it (Duncan) 

hmaieaentU ( Vautfian) 

M]Uafrr4p<«iui« (Vaughan) ... . 


X 
X 
X 
X 
X 
X 
X 
X 

X 

X 

X 
X 
X 













These three ''species" may be 
redooed to one. 


1 




::::;:.::::::: 


X 
X 










Ofifftitota (Duncan) 








eyoMAet (Duncan) 


















Troekotmiiia inaignU Duncan -f 


ProUthmot (^) new species 1 




i 


T. or^uia Duncan, nol BeoM. 
TrochosmiUa mbeurvota Duncan, 


new species 2 




1 


pLW,fi«.l,aotB«MB, 
Troehoamuiu tvbcurwUt Duncan, 


new species 3 




i 


pl.l9,fig.la,fM<B6ittS. 


new species 4 




i 




MtUtkmot (?) new species V, .'. . . .\ . . 








DendToeUcantahrifiensitVaxjg^'. 




. 


X 




iictinocit new species T 


X 
X 








MuUkolumnattnea cpaihIformU (Dun- 


X 




can) 




Aetinaeii roUd Duncan, noi 


new species 2 


X 




i 


Reuss. 
1*orHet rsfiiosa Duncan, nof 






Catulk). 



The following names in Duncan's list of St. Bartholomew corals 
are dropped, because the specimens on which he based his deter^ 
minations could not be found: 

1 Duncan, P. M., and Wall, O. P., A notice of the geology of Jamaica, especially with referenoe'to the 
district of Clarendon; yith descriptions of the Cretaceous, Eocene, and Miocene oorals of the Island, Geol; 
Soc. lyondon Quart. Joum., vol. 21, pp. 1-15, pis. 1,2, IMS (the descriptions of the oorals are by Duncan), 
Duncan, P. M., On the fossil oorals (Hadreporaria) of the West Indian Islands, (Hoi. Soc London Quart. 
Joum., voL 24. pp. 0-33, pis. 1-2, 1867. 



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OBOLOOY AKD PALSOKTOLOeT OF IHB OANAL ZOJTB. 196 

Stylophora distans (Leymerie). 
conferta^UBB. 
iuberosa Reuss. 
granulata Duncan. 

Stephanoeomia elegans (Leymerie). 

Agtrocoema muliigranosa Reuss. 
nmiosa (Sowwby). 

PlocophyUia caUculaia (Catullo). 

Solenastraea cohinmans Reuss. 

The revised list of the St. Bartholomew coral-fauna contains 33 
species, two of which may be referred to the synonymy, but a few 
species may be added from the collection I made, the study of which 
is not quite complete. I have described and have had figures made 
of all the species in the Oeve collection. I hope soon to add descrip- 
tions of the specimens I obtained and then to publish a full account 
of the fauna. 

I seriously doubt the Catadupa corals being Eocene; it seems more 
probable that they are Cretaceous. The species I described as 
TroehomdUa MUi is probably a f ungid coral. The Richmond '' beds '^ 
of Jamaica contain two species, one of which is found in the St. Bar- 
thdomew limestone. The Cambridge ''beds" contaui three species, 
two of which also occur in the St. Bartholomew limestone. The 
correlation of the Richmond and Camlmdge formations of Jamaica 
with the St. Bartholomew limestone, seems to be well founded. 

JACKSON FORMATION AND OCALA LIMESTONE. 

The corals of the upper Eocene Jackson formation in the Qulf States 
are described in monograph cited below.^ The species are as follows : 
FTabeUum cuneifarme var. wailesi Conrad. 
AldrichieUa ' elegans (Vaughan). 
Turbinolia pharetra Lea. 
Troehocyatkus lunuUtifarrnM (Conrad). 

var. montgiynurienm Vaughan. 
OaryaphyUia daUi Vaughn. 
PanumUia ludavidana Vaughan. 
Arehohdia bumsi (Vau^^n).* 
Astrofigia expanaa Vaughan. 

ludoviciana Vaughan. 

harrisi Vau^^n.^ 
PUUycoenia jacksonensis Vaughan. 
BalanophyUia vrarata (Conrad). 

1 VmgbH^ T. W., TiM Soensaiid lower OliffOiHMooml iMiitt of tteUn^ 
MoB.10,pp.9aS»Mikli.,lfl00. 6Meq>MioU7p.aO. 
tchaneed ftXMii iiUrfaMs. 



«Na; 



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196 BXTLLSTIK 103, UNITED SIAIBS KATIOKAL 1CU8BUM. 

Endopachys madurii (Lea). 

var. triangulare Conrad. 
shaleri Vaughan.^ 
mintUum Vaughan. 
A comparison of this list with the one of the St. Bartholomew 
and Jamaican corals reveals nothing in common; but I believe it can 
be made clear that the two faimas are of nearly the same age. That 
the Jackson formation in Mississippi and Loiusiana is i shallow-water 
deposit is indicated by the nature of the sediments, the growth of 
specimens of Astrangia on rounded, somewhat indurated balls of 
sand, such as are common along some beaches, the presence of 
oyster shells, etc. The striking difference between the Jackson and 
St. Bartholomew coral faunas is due neither to great difference in 
geologic age nor to difference in the depth of water in which the 
faunas lived, but it is due to difference in the temperature of the 
water. The St. Bartholomew is a tropical fauna; the Jackson is a 
temperate fauna. 

The correlation of the St. Bartholomew limestone, the Richmond 
and Cambridge formations of Jamaica, and the Brito formation of 
Nicaragua with the Jackson formation of the Gulf States has been 
made possible by the work of C. W. Cooke and J. A. Ciiahman. 
Cooke shows in the paper cited in the footnote ' that the Ocala 
limestone of soutiiiem (Georgia and Florida is of Jackson age; and in 
more recent papers he ' describes the stratigraphic occiu*rence, and 
J. A. Cushman^ describes the species of the orbitoid genus of foram- 
inifera Otihophragmina from the Ocala limestone in southern Geoigia 
and Florida. The following is a list of the species: 
OrO^ophragminaJlintensis Cushman. 
floridana Cushman. 
americana Cushman, st. 
mariannenais Cushman, st. 
mariannensis var. papiUata Cushman, «i. 
gecrgiana Cushman, st. 
vaughani Cushman, sU 
Those species whose names are followed by ''«i." are stellately 
marked or are stellate in form. The Ocala limestone is a shoal-water 
deposit, laid down in a sea having a tropical temperature.' One of 
the results of my collecting in St. Bartholomew was to find in the 
St. Bartholomew limestone a stellate species of Orihophragmina, 

> Name added. 

« Cooke, C. W., The age of the Ooala limestone, U. 8. Geol. Surrey Prof. Pap. «5-I, pp. 107-U7, lOlA. 

• Coolce, C. W.| The stratigraphic positloii and fjumal associates of the orbitoid foraminllBri of the gams 
Orikopkragmina from Oeorsta and Florida, U. 8. Qeol. Surrey Prof. P^. IOM>, pp. 10»-1U, IttT. 

4 Coshnum, J. X,, Orbitoid foramlniliera of the genus OHhop kr of mi n B from Georgia and Florida, U. 6. 
Geol. Survey Prof. Pap. 106-Q, pp. 115-124, pis. 4(M4. 

• Vaughan, T. W., A contribution to the geologic history of the Floridian Plateau, Carnegie Inst. Wash- 
ington Pub. 133, pp. 150-153, 1910. 



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GBOLOQY AND PALBOKTOLOGY OF IHS OAKAL ZOKB. 197 

nearly related to 0. mariannensis Cushman, and a second species of 
Orihophragmina that is of lenticular form. I also collected two 
species of Nummvlites and one species of Lepidocytlina in St. Bar- 
tholomew. Lepidocyclina occurs in Georgia as far down strati- 
graphically as a horizon about the middle of the Jackson formation, 
and apparently as low as the base of the formation. The presence of 
a species of Orihophragmina so similar to 0. mariannensis seems to 
wariant the correlation of the St. Bartholomew limestone with the 
upper part of the Ocala limestone of Florida and Geoigia; and there- 
fore witiii the Jackson formation in Mississippi and in the States 
farther westward. 

Regarding the Brito formation of Nicaragua » it must be recognized 
that a single poor specimen pf coral furnishes slim evidence on which 
to base a correlation. Doctor Cushman submits the following state- 
ment regarding the foraminifera from the Brito formation: 

Ab to the Brito matoiAl, two lots especiaUy are of interest. No. 6411 ''coast about 
2 m. s. e. of Brito Harbor" marked ''Ool. fos. 1. s.'' has abundant orbitoids with a 
beautifully ornamented exterior which without the confirmatory evidence of sections 
Beem to be clearly Orihophragmina of a group not so far represented in the material 
studied. From No. 6408 two miles n. w. of Brito Harbor, however, there is more 
evidence. The material is very different and contains specimens which in accidental 
section show definite chambeiB of Orthopkragmima of a different group. This does not 
however suggest either of the species from St. Bartholomew. Associated with it is a 
species of the flattened, broadly spiral form of nummulites. In the St. Bartholomew 
mat^ial there is such a form but of a species very much laiger. 

Now there is on the other hand a closer resemblance, that is to the lowest material 
oi the Flint River collections. The Brito species of Orihophragmina is similar so far 
as I have made out to the one I have called 0. JtinUntU, ICoreover it is associated 
at Brito as along the Flint River with this broadly spiral, flattened form of nummu- 
lite. The specimens of nummulite from the two localities are very close in form 
and size and only differ in minute details. They may not be specifically identical 
in final analysis but are very dose. 

The statement by Doctor Cushman seems conclusive. 
A horizon very nearly the same is recognizable in Colombia as the 
following quotation from Doctor Cushman shows: 

Now, as to the specimen from one league west of Arroyo Hondo, Bolivar, Republic 
of Colombia. There is an association of NummtUites and steUate orbitoids which very 
decidedly suggests Eocene. While I can not definitely make out the equatorial 
chambers, the stellate form is very apparent in several specimens, and I should say 
q>ecifically different from any of the species of Orthophragmina described in my 
paper from Georgia and Florida; in fact, they represent a very different group, I 
think, but are undoubtedly Orthophragmina. 

Ek>cene deposits of the same or nearly same horizon as the St. 
Bartholomew limestone are widely distributed in Cuba, as is indi- 
cated by species of Orthophragmina and a number of echinoid species 
that also occur in St. Bartholomew. 



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198 BULLExnr los, xtbtitid mATES KAnoKAL uwauu. 

CONCLUDING REMARKS ON THS EOCENE. 

From the foregoing discussion it is clear that marine upper Eocene 
formations are wfdely distributed in the southern United States, the 
West Indies, Central America, and northern South America, and 
that the Atlantic and Pacific Oceans were connected at that time. 
One of the areas in which there was such a connection was across 
the present site of eastern Nicaragua. 

Haug, I believe, correctly correlated the Jackson of Mississippi 
and other Gulf States with the Bartonian-Ludian (Priabonian) of 
Europe.* 

Attention should be directed to a statement by Opp^iheim ' in 
which he suggests that the St. Bartholomew coral-faima might be 
the equivalwit of the Priabona formation. The sequence I am 
giving three of the important American horizons precisely paral- 
lels Oppenheim's order, as expressed on page 13 of his work cited. 
It is as follows: 

Oligocene: 

Middle (Stampian <=RupeUan ==» Antiguan). 
Lower (Sannoisian » Lattorfian » Vicksburgian) . 

Eocene: 

Upper (Priabonian =* Ludian» Jacksonian = horizon of St. 
Bartholomew limestone, etc.)^ 

Oligocbnb. 
lower oligocene, 

The lower OUgocene corals of the United States have been de- 
scribed by me.' 

Dr. C. W. Cooke, in a paper recently pubUshed, subdivides the 
Vicksburg group in Mississippi, Alabama, and Florida as follows: 

Subdivisions of the Vic^btarg group in Mississippi, AUd>amaf and Florida. 

Mississippi. AlBlMnuu Floclda. 

Bryam calcareous marl. 



Qlendon limestone member. 



Mint Spring ! "Chlmiiey Rock" fades 

calcareous marl 

member. \ 



Forest mu sand Red BlulT clay 

(Western Hissfssippi). (Eastern Mississippi). 



1 Haug, 6mile, Traits de g^logie, vol. 2, p. 1628, 19U. 

* Oppenheim, P., Die Priabonaschicbten and ibre Fauna, Palaeontographica, vol. 47, pp. 348, 21 pis, 
1001. 

• Vaogban, T. W., Tbe Eocene and lower Oligocene coral faunas of the United States, U. 8. QeoL Survey 
Mon. 89, pp. 288, pis. 24, 1000. See espedaUy p. 80. 



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QBOLOGY AKD PALEONTOLOOY 01* THB OAKAL ZOHS. 



199 



The "coral limestone/' formeiiy referred to tke top of the Vicfa- 
burg group, as will be shown on subsequent pages, is, in my opinion, 
equivalent to the basal part of the Chattahoochee fOTmation. The 
fcdlowing IB a list of the species of corals at pres^it known from the 
Vicksburg group : 

Fossil corals from the Vteksbvrg group. 



Name. 



Byram 
calcareous 



FMdlum magnoeottatumYaagJiBn. 

rhomboideum Vaughan ... 

Twbinolia intiffnifiea Vaiuhaii 

Steriphonofnekua puleker vaughan. . 
ArckoktUa tu^ecta (Vaugltian) 

riaubuTgefuit (Conrad) . . . 

missUtmpieiutt (Conrad) . 

hoTTisi (vaughan) . 



a2drieAi(Vauehan)... 
** ' (puncan). 



Antifuattrea edhtiota (pu 

Saianophpllia eloiijato vaughan. 

eauiuera (Conrad) 

eatOi/era var.mukigranotaVuai^baji... 
DtniropkffUia new spwAM 



X 



limestone. 



Mint 

Spring 

limeRtone|ca1careou8 

marl 

member 



Qlendon 
limeittonc 
member. 



Bed 

Bluff 
day. 



X 
X 
X 
X 
X 
X 



X 
X 



X 
X 



This fauna is different from any now known in iJiie West Indies or 
Central America. It lived upder conditions closely similar to those 
under which the Jackson fauna of the same area lived. It is impor* 
tant to note that AnUffuaairea ceUvloaay a species very abundant in 
the middle and sparingly present in the upper Oligocene, occurs in 
the uppermost beds of the Vicksburg group. The Oligocene coral reef 
represented by the ''coral limestone'' at Salt Moimtain, . Alabama, 
and at Bainbridge, Georgia, overUes the Vicksburg group, which can 
with considerable assurance be correlated with the lower (Migocene 
(Lattorfian) of Veneto and elsewhere in Europe. The greatly- 
developed Oligocene coral reefs of Antigua are to be correlated with 
the reefs of Bainbridge. They are therefore stratigraphically higher 
than the Vicksburg group and are of middle Oligocene (Rupelian « 
Stampian) age. 

IHDDLE OLIGOCENE. 

AKnaUA FOEMATION.» 

The following list of species is based on a revision of Duncan's 
work on the Antigua corals,' after a study of his types in the collec- 

1 Name proposed by J. W. Spencer in his paper entitled On the geological and physical development 
of Antigua, Oeol. Soo. London Quart. Joum., vol. 57, pp. 49(M98, 1901. See also. Brown, Amos P., 
Notes on the geology of the Island of Antigua, Acad. Nat. Sci. Phila. Proc. for 1013, pp. 584-616, pis. 18-90, 
1913. Vaughan, T. W., pi^wrs referred to in footnote on page 103; and Memorandum on the geology and 
gnmndwaters of Antigua, B. W. I., Imperial Dep't of Agriculture West India Bull., voL 14, No. 4, 5 pp., 
1915. 

* Duncan, P. M., On the fossil oorab of the West Indian Islands, Part 1, Oeol. Soc. London Quart. Joum- 
vol. 19, pp. 40»-458, pis. 13-16, 1863; Part 4. Idem., vol. 24, pp. 9-33, pis. 1, 2, 1867. 

3714^—19— Bull. 103 2 



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200 



BULLETIN 103, UNITED STATES NATIONAL MUSEUM« 



tions of Geological Society of London and the British Museum 
(Natural History), and principally the collection made by myself 
which contains 60 species. It seems that I failed to find 7 of the 
species reported by Duncan; and apparentiy Mr. Robert T. Hill and 
Dr. J. W. Spencer each obtained one species that I did not collect. 
I feel a Htde doubtful about two or three of Duncan's types having 
really come from Antigua. Each species whose name is preceded 
by an asterisk * is considered in the systematic part of this paper. 

Fossil corals from the Antigua formation. 



Name. 



*8tifU)pkon ponderota Vauf ban. . . . 

newspecfesl 

new species 2. 

PociUopora tentU» Duncan 

new species 

Jiadrepora new species 

*Styloeoenia pumpeUpi (Vaughan) . 



new species 

*Attr6coenia mtafOanamefuit Vaughan. 
^decaiuren^ Vaughan 

^portoricentit Vaughan 

new species 

Atteroimilia exarata Duncan var 

Genus indet. new species 

SupkpUia new species 

Antiuia new species 

Leptomutta newspecies 

Qenuslndet. newsnedes 

Cladocora recrescent Lonsd ale. 



*OThieeUa antiUarum (Duncan). 
*coittta (Donoan) .... 



*in8igfUi (Duncan) 

*intermedta (Duncan) . . 

*Antigu<utrea celluUna (Duncan) . 



■hrar. curvoto (Duncan). . . . 
♦var. silecentis Vaughan . . 
IHpMhieattraia iiion^lor (Duncan) Dun- 
can. 

*Fa0tonMciofMltf< Vaughan , 

*F\ivUa polygtmaXi* (Duncan) 

new species t , 

Lamelltutraea tmythi Duncan 

Oennsindet. newspecies 

Ooniastrea reussi (Dimcan) , 

*M€eandra an/i^u^fwif Vaughan , 

deru-tlephanti* (Duncan) . . . , 

*Leptoria tpenceri Vaughan , 

* Mankina wtilougkbieMis Vaughan 



*Pironattraea antiguetuU Vaughan . 

Pawna new species 

Leptoseris new species 1 

new species 2. 

Haloteris new species.*. 

*8ider(utrea eonferta (Duncan) 



*Cifaihofnori^a hiUi Vaughan 

*brovmi Vaughan 

*temVaughan 

*9pimdena Vaughan . . . 
*antiffti,en9is (Duncan) . 



*Unuit (Duncan) . 



Distribution outside Antigua. 



Salt Ht., Ala . 



Balnbridgb. Oa.. 



Cuba, Panama 

Cuba. Bainbi'idge, Ga. 



Porto Rico 

Santo Domingo. 



Chattahoochee formation, up- 
per part, Oa. 

Montserrat 

Porto Rico, Canal Zone, An- 
guHla. 

Arube. 



Porto RJco, Cuba, Fla., Ga., 
Miss., Ilex., Anguflla, Arube. 



Ga., Fla., Mex. 



Bahibrldge, Ga.. 



Pj 
Cuba! 



Cuba.. 



Porto Rico, Canal Zone, An- 
guilla. 



Porto Rico, Cuba, Mex. 



Porto Rico, Cuba. 



Nomendatorial notes. 



Stjfloeoenia Idbato-rotundatm 
Duncan, noi M. Edwards 
and Hamie. 



Astrocoenia ornata Duncan, 
not M. Edwards and Haime. 



Heliattnta antiUarum Duncan. 
HelUutraea toUata (Dunoao) 

Duncan. 
Helkulnea inthnit Duncan. 
Heliattraea radiata var. filler* 

nedia (Duncan). 
Hdiaatraea eeUtUosa (Duncan) 

Duncan -h ItutnMatwhinmtm 

Duncan. 



Stephanocoenia revs9i Duncan. 
OouoriadentelepktttUi* Duncan. 

Ifaeandrina species Duncan. 
Coeloria UbvnntMfmriUa Don- 
can, noi Lmnaeus. 



Iwttnua conftrta Duncan. 



Heliattraea antignentiM (Don- 
can) Duncan -|- Atfnria 
ajjlnit Duncan -f Aatmia 
antiguenait Duncan. 

HeUattraea leimif (Duncan) 
Duncan. 



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QEQLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 201 

Foml corals from the Antigua formation — Continued. 



Name. 



Distribntton outside Antigoa. 



Nomeiidatorial notes. 



^IHpUmatna era$9oiameUata (Duncan) . 



*Far. tBOfiM/Scs (Duncan). . . 

♦var. nugenti (Duncan) 

l>tfiitfropj|fraia new species 

^Acrojwra pawunensU Vaughan 

new species 1 

*9aluaen»i» Vwighan 

new species 2 

^Attreoporaanti^uenti* Vaughan 

^AtaHQeii alabamknti* Vwighan 

new species 1 

new species 3. 

OoRlopora new species 1 

new species 3. 

*regularis (Duncan) 



Porto Rico, Cuba, Ga., Canal 
Zone. 



Porto Rioo, Cul>a, Qa. 



Canal Zone' . 
(}aiialZone.. 



Canal Zone 

8aU Mtl Ala.: Bainbridge, Qa. 
Batnbridge, Qa , 



*var. mkroteopka (Duncan). 
newmeoiesS 

♦ctow Vaughan. 



^IwrMMfiM Vaugban. 
" - • " ugfian... 



*aucadensi9 Vaugi 

new species 4 

new species 5 

Omtkopora (?) ttmU$ (Duncan). 



AnguHliL CkdsI Zone. . 

Porto Rico 

Anguilla, Canal Zone.. 



A Iveopcra new speciesl ., . 
newspeciesa.. 



HeUattraea cnuMiameUtUa 
(Duncan) Duncan + vara. 
magnttica, pulckeUa, and »»- 

+ var. minor Duncan. 



Alveopcra daedalaea yar.regur 
laria Duncan. 



SUphanocoenia tenuis (Dun* 
can) + Rhodaraea irregularit 
Duncan. 



Three of the species recorde(i by Duncan from Antigua, in my 
opinion, are incorrectly identified and their names are dropped 
from the list. They are as follows: 

Favoidea junghuhni Reuss, according to Duncan. 

Heliastrdea harbademis Duncan. 

SolenoBtrdea turonensis (Michelin), according to Dimcan. 

Another species, Astraea megaiaxona Duncan, is based on imiden- 
tifiable material, and its name is also (h*opped. The total number 
of recorded species from Antigua, therefore, is 69, and 5 varieties are 
recognized. Of the 33 species indicated as new, descriptions of 8 
have been written and descriptions of 26 remain to be written at 
the time of making out the preceding table. 

The number of species, 69, recognized is interesting for compari- 
son with the number recorded for areas in which Hving reefs occiur. 
Von Marenzeller ^ records 71 species from the Red Sea in his report 
on the Pola expedition corals Bedot ' records a total of 74 species 
4-5 varieties from Amboina — a number that should be reduced by 
about 4, because of the reference of some names to the synonymy of 
other species listed, leaving the number of valid species at about 70. 
In my paper on the shoal-water corals from Murray Island, Austra- 
lia, I Ibt 63 species from Murray Island and its vicinity in water 
not exceeding 18 fathoms deep, and report 51 species from Cocos- 

I VoD Marenzeller, B., Riflkorallan, Bzped. 8. M. Schifl Polo in das Rote Meer, Zool. Ergeb. », K. K. 
▲ted. WIM. Wlen, Hat.-Naturwlss Cl., yoI. 80, pp. 3S-07, pis. 1-39, 1906. 
* B«dot, M., Madr^poraries d' Amboine, Rev. soisse de Zool., vol. 16, pp. 14d-292, pis. 6-60, 1907. 



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202 



BULLETIN 103y UNITED STATES NATIONAL MUSEUM. 



Keeling Islands.^ It is known that at least a few more than 51 
species occur in these islands. Outside the main coraln^ef zone 
the number of species is smaller. For instance, there are only 43 
supposedly valid species recorded from water between and 25 
fathoms deep in the Hawaiian Islands and Laysan.' It is not cer- 
tain that 3 of the species included in the number 43 were obtained 
in the Hawaiian Islands. The usual number of species obtained iB 
Florida or the West Indies, in water lees than 25 fathoms deep, 
where conditions are favorable for coral growth is about 35. There 
were on the Antiguan reef as many species of corals as are at present 
usual for one island or a small group of islands in the Indo-Pacific, 
and about twice as many species as are usual on a Uving West Indian 
reef. 

The reason for referring the Antigua reefs to a horizon above the 
lower (Lattorfian) Ohgocene is given on page 199. The following 
list of middle (RupeUan) Ohgocene genera is taken from Fabiani,* 
but it is considerably revised and needs further revision: 



Middle Oligocene (Rupelian) genera of corals in Veneio, 



*Stylophora. 

*Stylohoenia. 

*Aslrocoenia. 

TrochosrrUlia,, 

Coelosmilia. 

Epismilia? 

PhylUmnilia? 

Parasmilia. 

*Euphyllia. 

Dichocoenia. 

Stylina7 

Ommia. 



MorUlivaultia'f 

Leptaxis. 

Astrangia. 

Holangia. 

Gombertangia. 

*Orbicella. 

Solenastrea. 

*Antigua»trea. 

Avlophyllia^ 

Rhabdophyllia, 

CalamopnyUia 

*Gon%astrea, 



*Hydnophora* 
^LeptomuMsa. 
Mycetophyllui. 
*Trocho9eri8. 

^Mesomorpha. 

Comoseris^ 

Myeetoseris. 

LeptophylUa? 

Stephanosmilia. 

ThanmasUria? 



Dimorphastrea^ 

Hyiiwphyuia. 

Astraeomorpha? 

AcrowKm. 

DenarfMci$, 

*Astreopora. 

*Aet%naei$. 

*G<miopora. 

*Porite$. 

*Alveopom, 



* Indicates that the genus is also found in the middle OUgooene of the West Indies or the aoutheastani 
United States. 

The generic characters of a number of the corals listed by 
Fabiani can not be ascertained without a restudy of authentically 
identified specimens in the Ught of modem systematic technique, 
which require that besides having an adequate knowledge of the 
morphology of the coral skeleton, the investigation shall proceed 
from a critical study of the type-species of the genera to be recog- 
nized to a similar critical study of the species to be generically iden- 
tified, and that due attention shall be paid to the rules of zoologic 
nomenclature as expressed in the International Code. I will point out 
in passing that there are in the United States National Museum 10 
specimens of the coral to which Reuss apphed the name CyathophyTUa 
annulata. It would be too great a diversion to give in this place a 
discussion of the literature on this species. This is a fungid coral, 

1 Vaughan, T. W., Some shoal-water corals from llnrraj Island (Australia), Coeoe-EeeliBg Is]aiuls,aiid 
Fanning Island, Carnegie Inst. Wadiington Pub. 318, see especially pp. 07-72, 1918. 

s Vaughan, T. W., Rooent Biadreporaria of tiie mtwalian Islands and Layian, U. S. Nat. Mas. BolL 
No. 59, pp. 32-34, 1007. [The list referred to has been sli^tly revised and the number redneed by 2aanm.l 

• Fabiani, R., U paleogene del Veneto^ R. Uniy. Padova Inst. Qeol. Mem., toI. 8, pp. 23»-381. 1916. 



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CffiOLOQY AKD PALEONTOLOGY OF THE OAKAL ZONE. 203 

with a synapticulate and perforatf^ wall at and just below the calicu* 
lar margin, the wall at lowet levels usually, but not inyariably^ 
becoming solid. In Fabiani's Ust this species, under the generic 
name Stephanosmilia (name proposed by Reuss in 1874, not Ste- 
phanoamiiia De Fromentel, 1862), comes between ParasimUa and 
PJocophyUia (a synonym of EuphytHa). I do not know what the 
systematic relations of Leptaais Reuss are. Reuss based the genus 
and the type-species, L. elUptica Reuss, on a single specimen from 
Monte Grumi and seems not to have obtained another from any- 
where. Until additional specimens of £. eUiptica have been critically 
studied, Leptaxis is not an identifiable genus. Although Dimcan 
considered Leptaxis a subgenus of AniiUiaj I think that it may be 
one of the simple fungid genera. The species referred to 10 genera, 
whose names are followed by a question mark," ?," should all be 
critically restudied. 

The names of the genera preceded by an asterisk, "*", in the 
foregoing table are also found in the middle Oligoc^xe of the West 
Indies or the southeastern United States. The following genera 
have closely related Bpecies: 

SfyUypTwra EuphyUia Leptomussa Adinaeis 

Stylocoenia OrhiceUa OyathcTnarpha Ooniopora 

Astrocoenia Antiguasirea Astreapara Alveopora 

I am not at all sure that some of the American middle Oligocene 
and the European Rupehau species are not identicaL 

Di^ Joseph A. Cushman has described the following species of 
Lepidoq/dina from the collection I made in Antigua (not yet pub- 
lished) : 

Lepidocydina gigas Cushman 

undulata Cushman 
undosa Cushman 
favosa Cushman 

L. undulata seems to be the largest known species of Lepicocydinay 
some specimens attaining a diameter of 100 mm. 

The calcareous algae, echinoids, MoUusca, and Bryozoa, as well as 
the Foraminifera of the Antigua formation will be described in a forth- 
coming volume to be published by the Carnegie Institution of Wash- 
ington. The Antigua formation must, in my opinion, be the type 
of the American middle Oligocene. 

P17INO FORMATION OF PORTO RICO.l 

The corals here listed were almost all collected by Mr. R. T. Hill. 
I have added the names of a few additional species collected by 
members of the New York Academy Porto Rico Siu^ey. 

1 For aooounts of tbe geologic relotioiifl of (taU fomution, tee Hill, R. T., Notes on the tomBt oonditloDS 
of Porto Rioe, U. 8. Dept. Agriculture Div. of Forestry Bull. No. 25, pp. 14, 15, 1880. Vaugtun, T. W. , 
we references in footnote on pp. 103,205. 



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204 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Foml corah from the Pepino formation. 

Aatrocoema portoricensis Vaughan, Antigua, and Canal Zone. 
OrbiceUa costaia (Duncan), Antigua, Anguilla, Canal Zone. 
Antiguas^ea ceUvlosa (Duncan), Antigua, Florida, Georgia., etc. 
Maemdra portoricensis Vaughan. 
Leptoseris portoricensis Vaughan. 
Pironastraea ar^uiUensis Vaughan, Anguilla. 
Siderastrea conferta (Duncan), Antigua, Canal Zone, Anguilla. 
Oyathomorplia antiguensis (Duncan), Antigua, Cuba, Mexico. 

tenuis (Duncan), Antigua, Cuba. 
Diploastrea crassolameUaia (Duncan), Antigua, Cuba, Qeoi^a. 
Astreopora portoricensis Vaughan. 
Goniopora portoricensis Vaughan, Antigua. 

Of the 12 species from the Pepino formation, 8 are known in the 
Antigua formation of Antigua. 

UmsfONK ABOTS OONdLOlinULTB ITKAB QVAMTlVAUO, OVBA. 

The geologic relations of the corals from the vicinity of Ouanta- 
namo will be described by Mr. O. E. Meinzer in a forthcoming report. 
The following is a list of the species: 

Fossil cordis from ths middle OUgocens^ OuantanamOf Cuba. 

PodUopora guantanamensis Vaughan. 

^strocoenia guantanamensis Vaughan, Antigua, Panama. 

decaiurensis Vaughan, Antigua, Georgia. • 

meinzeri Vaughan. 
ArUiguastrea ceUuhsa (Duncan), Antigua, Porto Rico, etc. 
Trochoseris meinzeri Vaughan, Panama. 
Pironastraea antiguensis Vaughan, Antigua. 
Oyathomorpha angmUensis Vaughan, Anguilla. 

arUiguensis (Duncan), Antigua, Porto Rico, etc. 
tenuis (Duncan), Antigua, Porto Rico, etc. 
Diploastrea crassolameUata (Duncan), Antigua, etc. 
Goniopora decaiurensis Vaughan, Georgia. 

Of the 12 species here listed 7 are also foimd in Antigua; of the 5 
remaining species 2 are at present known from only one locality, 2 
occur elsewhere in association with a fauna of the same facies as 
that of Antigua, while 1 occurs in the base of the Anguilla formation. 

Limestoney Rio CanapUy Manasasas trail, Cuha, 

The following species were collected by Dr. Arthur C. Spencer: 
Leptoria spenceri Vaughan, Antigua. 
CyathomorpJia tenuis (Dimcan), Antigua. 
Diploastrea crassohmeUata (Duncan) Antigua. 
The first and second species of the above list were obtained at sta- 
tion No. 3473 of the U. S. N. M. record of localities for Cenozoic in- 



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(ZBOLOGY AJa> PALEOKTOLOOY OF THE OAKAL ZOiNB. 205 

vertebrate coUections. Specimens of (MJwphragmirha were obtained 
at tbe sfune station and indicate upper Ek)cene or lower Oligocene as 
the age of the rock. This matt^er will be further discussed in the 
forthcoming report on West Indian paleontology. 

BASAL PART OF CHATTAHOOCHEE FOBMATION IN QEORQIA.^ 

The localities at which the specimens of fossil corals were obtained 
are at Blue or Russell Springs on Flint River about 4 miles below 
Bainbridge, and at other localities along Flint River to Hale's Land- 
ing, about 7 mUes below Bainbridge. The corals are most embedded 
in or weathered out of chert which was once a coral-reef limestone 
that was formed on the subaerially eroded surface of the Eocene 
Ocala limestone after submergence. Dr. W. H. Dall in a recently 
published paper * appears to correlate this bed with the OrthoAilax 
pugnax zone of Tampa, Florida, and states that I concur in that 
opinion. Although the chert forming the base of the Chattahoochee 
formation in the vicinity of Bainbridge is faimaUy nef^ly related to 
the "silex" bed of the Tampa formation, in my opinion they are not 
of the same age, the ''silex" bed being geologically j'oimger. The 
coral faxmas are not the same, and there is at least a species of one 
genus at Tampa of stratigraphically later afiBnities than any species 
in the vicinity of Bainbridge. 

The following are the species from near Bainbridge mentioned in 
this paper: 

Fossil corals from basal part of Chattahoochee formation near Bainbridge^ Georgia, 

Siylophora minvtismna Vaughan. 

Stylocoenia pumpeUyi (Vaughan) Vaughan, Antigua. 

Astrocoenia decaturensis Vaughan, Antigua, Cuba. 

OrhiceUa bainbridgensis Vaughan, Santo Domingo ?, Porto Rico. 

Antigua^rea ceUulosa (Duncan), Antigua, etc., Tampa. 

YaT.'sUecensis Vaughan, Antigua, etc. 
Fcmtes polygonalis (Duncan) var., Antigua. 
Siderastrea sUecensis Vaughan, Tampa; Alum Bluff formation. 
Diploastrea crasaolameUata (Dxmcan), Antigua, etc. 

var. viagnifica (Dimcan), Antigua, etc. 
Astreopora antiguensis Vaughan, Antigua. 
Actinada alabamiensis (Vaughan), Antigua; Salt Moimtain, Ala. 
Ooniopora decaiurensiB Vaughan, Cuba. 

1 The more important references to the literature are as follows: 

Vaoi^ian, T. W., A Tertiary coral reef near Bainbridge, Georgia, Sdenoe, n, s., vol. 12, pp. 873-S76, 1900; 
BatBbridfi and vioiiitty in Prettminary report on the Coastal Plain ol Georgia by O. Veateh and L. W. 
Stephenson, prepared under the direction of T. W. Vaughan, Geol. Survey of Oa. Bull. 20, pp. 328"3389 
1911; Tbe reef coral fauna of Carrizo Creek, Imperial County, California, and its signifjcanne, U. S. Q^, 
Bnnrsy Prof. Pl^)- W-T, pp. 3(0^864, 1917. 

Cooka, G. W., Age of the Ocala limestone, U. 8. Geol. Survey Prof. Pap. 95-1, pp. 107-U7, 1915. 

* A ooBtoilNition to tbe invertebrate fauna of the Oligocene beds of Flint River, Georgia* Proc U. S. Nat* 
Ms., vol. 51, pp. 487-524, plates 88-88, 1916. 



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206 BULLETIK lOe, JJTSTLTED STATES KAHOKAL MUSEUM. 

Of the 13 species and varietiea listed aboye, 9 are common to 
Antigaa, and Goniopara decatwrenaia occurs in Cuba in association 
with specaes of corak abundant in An%ua; of the 3 remaining 
species, Stylophara miniUUsima has so far been poeitiyely identified 
only at Bainbridge, but it is very near a species common in Antigua; 
2 of the 13 forms are known from the "silex" bed of Tampa. The 
coral fauna near Bainbridge is a moderately rich one. In addition 
to those listed there are species of Styhpharaf Astroeoenia, AntiUiaf, 
Aetrangia or Rktzangia, Mes&morphn, Agtreop(mi, ActinaciSf Oonuh 
poraj and Alowpora, and of a few genera not yet positively ideiltified. 
There are between 25 and 30 species, of which only 4 or 5 are com- 
mon to* the Tampa coral f aima. 

It should be stated here that casts of 'a species of Pecteuy which 
appears to P. suwaneenais Dall, occur at station 3381 in the matrix 
with Diplodstrea craaaolameUcUa, which may therefore be of upper 
Eocene as well as of Oligocene age, or I may not hare discriminated 
closely enough between species. 

"CORAL LDCBSTOHB" OF SALT MOUNTAIN, ALABAXA.l 

I described in the monograph referred to in the footnote two 
species, as follows: 

Siylophora ponderosa Vaughan, Antigua. 

Adinads (ddbamienms (Vaughan), Antigua; Flint River, Georgia. 

I long surmised that the. ''coral limestone'' of Salt Mountain 
really represented the basal part of the Chattahoochee formation, 
but only recently did I obtain evidence that this limestone is the 
stratigraphic correlative of the Antigua formation and of the coral 
reef horizon near Bainbridge. 

SAN BATASL FOBICATION OF EASTERN MEZIOO.< 

The formation from which the fossil corals were obtained was 
first designated by Mr. Dumble ''San Fernando beds/' a name long 
in use for a Tertiary formation in the Island of Trinidad. He has 
recently changed the name to San Rafael. It is an important forma- 
tion in eastern Tamaulipas, Mexico. Several of the corals are not 
well enough preserved for purposes of identification. The following 
is a Ust : 

Ardiguasirea ceUvlosa (Duncan), Antigua, etc. 

var. dlecensis Vaughan, Antigua, etc. 

Fcmtes mezicana Vaughau. 

Maeandra dumhlei Vaughan. 

> For a descripth)zi of the geologic relations, see Vsoghan, T. W., Eocene and lower ONgooene oocal 
fatmas of the United States, U. 8. Geological Survey Mon. 89, pp. 80, 81, 1900. 

• The principal literatore is as foUows: 

Domble, E. T., Some events in the Eocene history of the present Coastal area of the Oidf of Mexioo in 
Texas and Mexioo, 7oum. Oeol., vol. 28, pp. 481-496, 1915 (see especially pp. 40^97); Tertiary deposits 
of northeastern Mexico, Califomia Acad. Sd. Proc., ser. 4, vol. 5, pp. 168-108, pis. 1^19, lfl5 (si 
dally pp. 18^192). 



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QSOLOOY AKD PALEONTOLOGY OF THE OAKAL ZONE. 207 

GyaJOtormorpha antigtLenMs (Duncan), Antigua, etc. 
Ooniopora species. Similar to Antiguan species. 
Altliough the identifiable species are few, it i^pears safe to cor- 
r^te the San Rafael formation with the Antigua fcmnation. 

TOMOat PANAMA. 

Doctor MacDonald obtained at this locality, station 6587, the fol- 
lowing species of corals: 

Astroeoenia gtuintanamerms Vaughan, Antigua, Cuba. 

Fwvia macdonaJdi Vaughan, Antigua. 

Maeandra ajUiguensis Vaughan, Antigua. 

Trockoseris meimeri Vaughan, Cuba. 

DipUxuirea erassolamettata (Duncan), Antigua, Cuba, etc. 

There can be no reasonable doubt that this is the same as the coral 
faima found in the Antigua formation. As the locality at which 
the specimens were ol>tained is on the Pacific coast of Panama, the 
evidence is conclusire that there was middle Oligocene connection 
between the Atlantic and the Pacific in that area. 

8BBB0 OOLOBABO, ABUBS. 

Three species were obtained at this locality,^ as follows: 
Orbicella iTrngnia (Duncan), Antigua. 
Antigtuuirea ceUutoaa (Dimcan), Antigua. 

Ooniopora species (the kind of casts to which Duncan appUed the 
name Alvepora daedaUa var. reguUms). 

This fauna is evidently the same as that of the Antigua formation. 

COMCLXJDINO KEMABK8 ON THB MIDDLB OUOOCKNE. 

The foregoing lists show that Antiguan middle Oligocene coral 
fauna is known in Porto Rico, Cuba, southern Georgia, southern 
Alabama, eastern Mexico, Panama, and Arube. That it also occurs 
in Santo Domingo is known from some of the specimens, Siderastrea 
conferia (Duncan) typical and a peculiar variety of Asterosmilia 
exarata (Duncan), both brought from Santo Domingo by Gabb. It 
is a key horizon in the American Oligocene. The Byram calcareous 
marl of Mississippi occurs either at its base or just below its base. 
It therefore overlies all the Vicksburgian lower OKgocene, with the 
possible exception of the uppermost member, and is stratigraphi- 
cally just below the ''silex bed'' of the Tampa formation. The 
correlation of the deposits containing this fauna with the KupeUan 
of Veneto has been made on pi^e 202. 

That there was middle Oligocene connection between the Atlantic 
and the Pacific was pointed out on this page in discussing the species 
from Tonosi, Panama. 

I Vsogtum, T. W., Some ftMsU corals ftom the elevated reefs of Caracao, Arube, and Bonaire, Qeolog. 
Bekte-Mitt. UMbd 8amml.» aer. 3, toL pp. 1-«1, 1901 (espedi^y pp. 11, U). 



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208 BULLETIN 106y UNITED STATES NATIONAL IfXTSBUM. 

UPPER OUQOCSNE. 

CUIJCBB4 FOEMATIOir. 

The Culebra formation and the base of the E}mperador limestone 
in the Canal Zone contain a few species that indicate close relation- 
ship with the Antiguan horizon, but on the whole the aflSnities are 
rather with the next higher fauna. Fossil corals were obtained in 
the Culebra formation at three stations, as follows: 

Station 5863, west side of Gaillard Cut, at station 1863 of the 
Canal Commission, between points opposite Curacha and Paraiso. 

Station 6020c, Las Cascades, Gaillard Cut, third bed from the 
bottom of the section. 

Station 6026, one and one-half miles south of Monte Lirio, on 
Panama Railroad (relocated line). 

The list of species is as follows: 

Species of corals from the Culebra formation. 



Name. 


8tatt<ii 
6868. 


SUtlOD 

W20e. 


StetiOD 
6020. 


^pen- 
dor Is. 


Anttgua. 


^^ 


StyU^pkcra imnfrat^frLt Vaughan - . r 


X 




X 
X 


X 
X 




X 


' " ^o<i*«W Vaughaii 


' 






Orbiedla eoHata (Puncan) . 


* 


XX 


X 
X 
X 
X 


X 


Siderattrea eon/erta (J )uncaii) 








X 


Attfeopcn fifUiffV4n*if Vn.ligha.n ^ , , 




X 






Oo^ioporti (tucad^iifif Vaughan . . , » . ^ . .... 





^ 




X 











Of the 6 species in the Culebra formation, 2 also occur in the 
Emperador limestone; 4 also occur in the Antigua formation; and 4 
also occur in the Anguilla formation. There is only one species, 
Astreopora antiguensiSf that is elsewhere known only from the 
Antigua horizon; while 2 species are at present known elsewhere 
only from the Anguilla horizon. These relations indicate, but do not 
prove, that the upper part of the Culebra formation, the part of the 
formation in which the corals were collected, is stratigraphically 
higher than the Antigua f ol*mation, and is, therefore, referable to the 
upper Oligocene. The foraminiferfd fauna, to be discussed on pages 
554, 555, 585, supplies stronger evidence in favor of considering the 
upper part of the Culebra as of upper OUgocene age. 



KMPBRADOR UmSTOini. 



The principal collections from the Emperador limestone were made 
by Doctor MacDonald and me at Station 6015 and 6016, in Empire 
village. Dr. Ralph Arnold subsequently made a small collection in 
Empire and obtained one species, PociUopora amoldi Vaughan, not 
collected by Doctor MacDonald and me. Doctor MacDonald and 
I also made a small collection at Station 60246, the upper bed at 
the lower end of the culvert where the Panama Railroad (relocated 
line) crosses Rio Agua Salud; and he subsequently obtained some 



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GEOLOGY AND PALEONTOLOGY OF THE OANAL ZONE. 



209 



very interesting specimens at station 6256, which is 1^ miles south 
of Miraflores. The following is a list of the species: 

Species of corals from fhf. Emperador limestone. 





Empire 
quarries. 


station 
6024b. 


Station 
6256. 


AngoOIa. 


Antigua. 


Slfla^piar^ intperaiorbf VaqghftD 


X 
X 
X 
X 
X 
X 


X 




X 




poiiaWAWt Vaugban 
















mitcdimaJ4i Vaagtian 


X 








M^am VanghaiT. 








PoctZlopora omoUi Yaiighan 










AMrocoerda, portoriceiuUiV&u^iUk 


X 






X 


OftfeeOa <mperator<« Vaagtian 


X 

X 
X 
X 
X 


X 


X 

X 




eanaliA Va^ighaxT. ...... . . ...... . 






Stfkmgia TpanamenaUfytM^mn 











GonioitrcaeaiiaZif Vaughan 








Pamma pamnnentU VMi^ian 










Aeropofu panamentia Vahg^iaii 


X 
X 






X 


ialvdjtnHt Vaug4ian t » 


X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 
X 






X 










OomiAjmmiimVf^^n 










pttnamtniU VAUf^bMi , 






X 
X 
X 
X 




Impcftlorit Vau^baa 
















i>i«EM< VftMgf^n 






X 


PoriUs dowiOei VAuAan 








iam^Mtmghfin.... 










2NiiiainCTw£i Vaughan 




X 






«VMflh«rif Vmigfaan 




X 


X 


iSynaraea) howel Vaudian 














X 


X 











Of 26 species from the Emperador limestone, 6 have been identified 
in the Antigua formation and 9 in the AnguOla formation, but it 
is probable that the number of species common to the Emperador 
limestone and the Anguilla formation will be somewhat increased. 
The Emperador limestone is of nearly the same horizon as the An- 
guiHa formation. Additional evidence favoring this opinion will be 
adduced on subsequent pages. 

ANGUILLA rOBMATION.l 

This name is proposed for the coralliferous limestone and argilla- 
ceous marls of Anguilla. The type-locality is on the south and west 
sides of Crocus Bay, where it is exposed to a thickness of about 200 
feet. The f aima has been monographically described, and the accoimt 
of it will be published in a forthcoming volume of the Carnegie Institu- 
tion of Washington. The following species of corals from it are 
considered in the present paper: 

> The principal ttterature Is as follows: 

OlBV, P. T.« On the geology ol the nortiieastern West India Islands, K. svenska Vet-Akad. HandL, 
ToL0,No.U,p.2^1872. 
Vaofhan, T. W., see references In footnote, p. 103. 



Digitized by VjOOQIC 



210 BULLETIN 103y X7KITED STATES NATIONAL MUSEUM. 

Speciei ofcoraUfrom the AnguxUa formation. 



Name. 



Culebra 
forma- 
tion. 



Empera- 

dorilme- 

stone. 



ADtigua. 



Other locaUttes. 



Stvlophora imperatoris Vaughan. 
Orbk^la impentori$ Vaogl 

co8tata (Dtmcan) 

conoiif vaughan 

A nttguagtrea eeUulosa (Duncan) 

Aoarkia atkffuittstuis Vau^ian 

Pironastraea anguiUensis Vaughan. . 



X 
X* 



X 
X 



Cuba. 



Sidtrastrea eonferta (Duncan)., 
^ " * I an^tUfn^ Vaughan. 

roTkorottghi Vaughan.. 



Ci/alhomorpha anguiUensis ^ 



X 
X* 



P. R.; Cuba; etc 



P. R. 
P.R. 



Ooniopora varuimtnsis Vaughan. 

trnp^rotorit Vaughan 

pana/i« Vaughan 

d«i;«f Vaughan 

ctucodfiMit Vaughan 

Porites anguiUensis Ywifiaua 

(Sjfnaraea) maedonaldi Vaughan. 



X 
X 
X 
X 



X 
X 



P.R.- Porto Rioo. 

Of the 17 species listed above, 4 are also found in the Culebra 
fonnation, 9 in the Emperador limestone, and 12 of the 17 in the 
combined Culebra and Emperador of the Canal Zone. In addition 
to the species here considered there are other species of Siylophara^ 
Stylocoenia, AntHlia, Cladocora, Maeandra, Ganioporay and Paritea. 
There are 9 or 10 species of Ooniopora. The total coral fauna in the 
collections available to me comprises about 28 species. 

The Anguilla formation is correlated with the E^mperador limestone 
for the following reasons: Heteroateginoides^ a new genus of orbitoidal 
foraminifera described by Doctor Cushman, is represented in the 
Anguilla formation by a species, also foxmd in Antigua, but very 
near a species that occurs in the Eknperador limestone. Although 
Heterosteginoides occurs in both Antigua and Anguilla, Lepidocydiiui, 
which is so abundant in Antigua, was not collected by me in Anguilla 
and is only sparingly present in the Emperador limestone. The 
identity of certain species of corals in the two formations has been 
shown. Echinolampds semiorhis Guppy is abundant in Anguilla (on 
the west side of Crocus Bay between 25 and 70 feet above sea level) 
and in the base of the Emperador limestone, Canal Zone. Orthavlax 
pugnax (Heilprin) was collected in the base of the Crocus Bay ex- 
posures. 



CUBAN LOCAUTIBS. 



OrbiceUa imperatoris Vaughan has been collected at the following 
locahties in Cuba: Station 3450, 4 miles north of Pinar del Rio; 
station 3451, one-half mile west of Cifinaga railroad station, near 
Habana; station 3566, Bejucal; station 7544, Rio Yateras, near 
Guantanamo. That the Anguilla horizon is widely extended in 
Cuba is shown by the distribution of the echinoids which will be 
considered in another place. 



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OEOLOGY AKD PALEONTOLOGY OF THE OANAL ZONE. 211 

f AMPA VOBMATIOF OF FLOBIDA. 

The corals from the ''silex" bed of the Tampa formation considered 
in this paper are as follows : 

OrbiceHa tampdensia Vaughan. 

var. sUecensis Vaughan. 

ArUiguastrea ceUvJosa (Duncan). 

Siderastrea sUecenais Vaughan, 

Siderastrea hiUshoroensis Vaughan occurs at about the same horizon 
as the ''silex" bed. 

The Tampa coral fauna has not been described in print, but I 
furnished Doctor Dall a list of my manuscript names of the species 
and it appeared in his monograph of the moUuscan fauna of the 
OrthauUix fugnax zone of the Oligocene of Tampa, Florida.* I have 
pointed out that OrbiceUa tampaenais var. sUecensis isee p. 391 of this 
paper) closely resembles some of the variants of 0. coatata from An- 
guilla and that the specimens identified as Sideraatrea ailecerhaia in 
which there are over 60 septa perhaps should be referred to 8. con- 
/erta (see p. 449). Besides the species mentioned, there are species 
representing the following genera: Stylopliora, AntiUiai, OaJaxea, 
Solenaatreaj Maeandra, SyzygophyUia ?, Endopachya, Acroporay Oonio- 
pora, Poriteaj and Aheopora, 

Two and perhaps three of the "silex" bed species of corals also 
occur at Bainbridge, but the faunas otherwise are not the same. 
Two of the species from Tampa are near living West Indian and 
Floridian species. These are Solenaatrea iampdenaia Vaughan, nomen 
nudum, which is near 8. hyadea (Dana) ; and Poritea wiUcaxi Vaughan, 
namen nudum, which has the septal arrangement of Poritea aaireoidea. 
The presence of such species with modem affinities seems to me to indi- 
cate a consid^ably younger age than that of the reefs near Bain- 
bridge. Furthermore LepidocydiTia is abundant in the reefs near 
Bainbridge, but has not yet been found at Tampa. Orihavlax pugnax 
occurs in the ^'silex'^ bed at Tampa, but it has not been foimd in the 
overlying limestone; the same species occurs in the base of the An- 
guilia formation, but I did not find it at higher levels. Dr. C. W. 
Cooke, who has monographically described the moUusca of the 
Anguilla formation, correlates it with the Tampa formation on the 
basis of similarity in their moUuscan faunas. The correlation of the 
Tampa formation is further discussed on pages 570, 571. 

CONCLUDOrO BXICARX8 ON THE UPfSB OUOOCKNS. 

That there was connection between the Atlantic and Pacific oceans 
during upper Oligocene time is shown by the continuity of both the 
Colebra formation and the Emperador limestone from the Atlantic 
to the Pacific slopes of the Isthmus. On the geologic map, plate 153, 

I VTS. Nat. Mas. BuU. 90, p. 18, 1915. 



Digitized by VjOOQIC 



212 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

the Emperador lunestone is represented as dipping below the Ghttim 
formation on the north side of the Isthmus, and it is exposed almost 
down to the sea level on the south side. The Culebra formation 
imderlies the Emperador limestone on both slopes, but it is not indi- 
cated on the map on the north slope of the Isthmus. 

MiOGSMB. 
BOW DEN MARL.1 

The point of departure in the consideration of the Miocene is the 
faima of the Bowden marl of Jamaica. The following is a revised 
list of the species: 

Placotrochus costatus Duncan. 

Sphenotrochu8 new species. 

Placocyaihus harretti Duncan. 

alveolus (Duncan.) 

Siylophora granuUUa Duncan. 

AsterosmUia profunda (Duncan). 
Tiilli Vaughan. 

Stephanocoenia intersepta (Esper), also living. 

AnLiUia waUi Duncan. 

Thysanua exceniricus Duncan. 
elegans Dimcan. 
new species. 

SyzygopJiyUia gregorii (Vaughan). 

Siderasirea siderea (Ellis and Solander), also living. 

Ooniopora new species. 

Porites harucoUensis Vaughan. 

Acropora new species. 

This fauna indicates somewhat deeper water than that in which 
the species mentioned on preceding pages Kved; but the presence of 
Siephinocoenia irUerseptaj Siderasirea siderea, Acropora new si>eci6S, 
a massive species of Ooniopora, and Porites baracodensis, furnish evi- 
dence in favor of the conclusion that the depth probably was not so 
much as 20 fathoms. The most striking feature of this list is that it 
contains the names of two species still living in the Caribbean region, 
in this respect differing from all the other faunas previously consid- 
ered in this paper. The Bowden not only marks the introduction of 
species that persist in the West Indian region, but as neither in 
Jamaica, Santo Domingo, nor Cuba, have species of Astrocoeniaf 
Siylocoenia, Leptomussa, Aniiguastrea, Fo/viies, Leptoria, Trochoseris, 
Leptoseris, Haioseris, Pironasiraea, Mesomorpha, Cyaikomorpha, 
Diploastrea, Astreopora, Aciinads, or Porites {Synaraea) been foimd 
in beds of the same age as or yoimger than the Bowden, these 

1 For an account of the stratlfraphic relations of the Bowden marl, seo Hill, R. T., The geology and 
pbyslcal geography of Jamaicfl, Mas. Comp. Zool. Ball., vol. 34, No. 1, pp. 290, wHh 85 plates, 78tf 
(especially pp. 82-88, 145-152). 



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QEOLOOY AND PALEONTOLOGY OF THE CANAL ZONE. 



218 



15 genera and one subgenus of middle and upper Oligocene corals 
apparently had become extinct in this region. [The genus Thysanus is 
present in the PUocene Caloosahatchee marl of Florida; and in Santo 
Domingo PlacocycUhuSj StylophorUy AntiUia, and SyzygophyUia occur 
at horizons above that of the Bowden marl, while the number of species 
now living increases. The Bowden marl marks an important change 
in the character of the coral faunas, a change from an older to a more 
recent facies. It therefore seems to me that the Bowden marl can 
not be considered of Oligocene age, and that it must be referred to 
the lower Miocene. 

SANTO DOMINGO. i 

With regard to the species reported by Duncan from Santo Do- 
mingo, it will be said that Dimcan does not describe the stratigraphy 
of Santo Domingo, but refers the specimens to the Nivajd shale, the 
superficial or tufaceous limestone, Posterero shale, Cerro Gordo 
shales, Esperanza shale, and ''the silt of the sandstone plain." The 
following is a list of the species recorded by him, the geologic forma- 
tion in which they were reported to be found, and the revised names 
with annotations: 

FostU corah reported by Duncan from Santo Domingo, 



Name used by Duncan. 


n 

oS 

? 

s 


•OS 

Ss 

if 

SB 


1 


i 

•g 

i 

1 


a- 


Kevised names and annota- 
tions. 


Flabellum exaratum Duncan 












Described from Vere, Jamaica; 


Flabellum new species 


+ 










genus doubtful; identifica- 
tion doubtful. 
Not determinable. 


PlaeotTOchus loTuiialei Duncan 










Placotrochus lonsdalH Dnrn^n. 


€kTatf>ttochus duodccim-costatus H. £d> 










From yellow shale of Angos- 


wards and Haime. 

TrocJiocyathus latfTo^pinoaus U. Edwards 
and Haime. 


+ 

+ 










tina, Santo Domingo; speci- 
men not determinable. 
Placocyc^hui new species. 










Parac^athut hmekeni (Dun- 


Placocyathtu barretti Duncan 




+ 






can). 
Plaeocyathus baTreUi Duncati; 


variabilh Duncan 


+ 






+ 


originally described from 
Bowden, Jamaica. 
Placoa/athus variabilis Duncan. 


costafus Duncan 


+ 
+ 

+ 

+ 








Piacocyaihus costatus Duncan. 


Pocillopora crasBoramom Duncan 

StflophoTo affinis Duncan 










Pocillopora crasioramosa Dun- 










can. 
Stylophora affini* Duncan. 


var. minor Duncan 










Stj/lophora viinor Duncan. 


StylophoTa afftnis Duncan var. 2 










Stvlophora new species; also 

from Cerro Gordo shales. 
Name discarded for Suito Do* 


mrisUlla M. Edwards and 








•f 


Haime. 
TTochocyathuM abnoTmalis Duncan ....,,, 


+ 
4. 








mingan species. 












lAsterosmilia abnormalii (Dun- 


comuta Duncan 










J can). 


exarata Duncan... 












A$Uro9milia exarata Duncan; 


SUphanoeomia iniertepta M. Edwards 

and Haime. 
AntillastTaea spOTtgiformis (Duncan) 

Duncan. 
Dkkoeoenia tuberota Dnneaii. 


....•• 


. — ... 






+ 
+ 


also in Antigua formation , 
Antigua. 










SUpharwcoenia inter*epia (E»- 

per). 


+ 


+ 












+ 





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214 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



Fossil corals reported by Duncan from Santo Domingo — Continued. 



Name used by Duacan. 



FUbeUum diMum Dunean. . 
AntaUa lorudaMa Duooan. . 

MIfffrflfti Pntiffftn 

C^phaslraea ooitata Duncan. 



PkjHIoeoenia iculpia M. Edwards and 
Haime var. tegtUa Dunoan. 

PkffUoeoenialimbata Duncan 

PUtitutraea ramai Dunoan 

BtUtttnea qfUniriea (Duncan) Doncan 
endotheeata (Dunoan) Dunoan 
brevk (Duncan) Duncan 

Plakutnea iUtant Dunoan 

^toftoM Duncan 

SoUmutnea vnkdtU U. Edwardi and 

Haime. 

SUphanoeoenia dendroidea U. Edwaids 
and Haime. 

Thptanut eorhtcuia Duncan 

TeliHaphifUia grandU Dimcan 

navkula Doneaa 

JUankina areoUUa (Linnaeus) 

JUaeandrina fiiogruna Lamarck 



sinuosisHma M. Edwards 
^ ^T^^ Haine. 



Lithophpllta affini$ (Dunoan) Dunoan.. 



AnMUt pondmta (K. Edwards and 
Haime) Dunoan. 
dentata Duncan 



Agarida agarkUa Lamarck. . . 

wndata Lamarck var.. 

Sidirattraea grandit Duncan. . . 



erenuUUa Blatnville var. an^ 
liZtorum Duncan. 



PorU49 eolkgniama Midhelin.. 
Alveopora fenettrata Dana... 



I'll 






a 



Revised names and azmota- 

ttODS. 



^XnmUidiiMi (Duncan). 

iintfttfolOabate Duncan. 
The type of this is from Bar- 



(ElUsa 

Santo DominjEan specimen 

is a species of SoUnattrm, 

Orhkella linibaia ( Duncan); alio 
reported flpora "yellow anale.'* 



Varietal forms of OibUaim cot- 

emo«a (^Linnaeus). 
OHrieella bTe9u (Dunoan). 
) Varietal fonns of Solenastrm 
} boumoni M. Edwards and 
] Haime. 

t A highlv fossilized specimen; 
I name discarded for the Santo 
I Domingan coral. 
! Locality not eiven: probably a 
, species of Aatroeoenki. 
T^fmmus eorMcula Duncan, 
rkyiaiitw ^rofMf if (Duncan). 
rJkfMiHw naofciOa (Duncan). 
I Ifaeandn areolatu (Linnaeus). 
I << Shale/' no other data on geo- 
logic relations; name dropped 
from list. 
The name proposed by Milne 
Edwards and Haime is a 
synonym of MkMmdn ttri- 
go9a (Dana); name dropped 
flrom list. 
JAwra affinit ( Duncan); may be 
the young of Mutta anguiMa 

type from Bowden, Jamaica. 

SjftfgopkglUa dentata (Dun- 
can). 

I Material poor, names dropped 

r fhunlist. 

Type from Jamaica is Sidermi' 
trea Merea (Ellis and So- 
lander). 

This seems to be a synonym of 
S. tiderea (Ellis and SoJan- 
der). 

Name dropped from list. 

Name dropped ftom list. 



It has appeared that perhaps two distmct geologic horizons w&te 
represented by these collections, one of which is the Nivajd shale and 
another which is represented by the superficial and tufaceous lime- 
stones and the silt of the sandstone plain. The revised list for the 
Nivajft shale is as follows: 

Revised list of species reported by Duncan from the Nivafi shale, 

Placotrochus lonadald Dimcan. 
Paracyaihus henekeni (Duncan). 
Placocyathus variabilis Duncan. 



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GBOLOOY AND PALBONTOLOQY OP THE OANAL ZOISTE. 215 

Placoeyaihus costatus Duncan. 

new species. 
Pocitlopara crasscramosa Duncan. 
Styhphora ajtnia Duncan. 
minor Duncan, 
new species. 
A8tera9milia abrwrmalis (Duncan). 

exarata Duncan. 
Dichocoenia tuberosa Duncan. 
ArUiUia dubia (Duncan). 
bilobata Duncan. 
Orhieetta limbaia (Dimcan). 
brevi8 (Duncan). 
cavernosa (Linnaeus). 
Thysanus corUcuia Duncan. 
grandis Duncan. 
navieula Duncan. 
Maeandra aredlata (Linnaeus). 
SyzygophyUia gregorii (Vaughaa). 
dentata (Duncan). 
A total of 23 species. 

The species from the superficial and tuf aceous limestones and the 
silt of the sandstone plain are as follows: 

Revistd list of species reported by Duncan from the superficial and tufateous limestones 
and the silt of the sandstone plain, 

*Placoeyathu8 varioMlis Duncan. 

Siephanocoenia inkrsepta (BJsper). 

*Dichoooenia tuberosa Duncan. 

OrbieeUa limbata (Duncan). 

"^OrbieeHa cavernosa (linnaeus). 

Solenastrea boumoni M. Edwards and Haime. 

Mussa affims (Duncan). 

Siderastrea siderea (E31is and Solander). 

A total of 8 species, of which 3, those preceded by an asterisk *, are 
also reported from the Nivajd shale; 6 of these species are either at 
present living in the West Indies or the fossil specimens are so similar 
to those of living species that specific discrimination is uncertain (see 
table on pp. 213, 214 for notes). One species, OrUceUa liwhata, is very 
similar to one of the growth forms of OrbieeUa anv/ularis. This leaves 
only one species, Placocyaih/as variahilis, that seems clearly to indicate 
an older Tertiary age. But it should be added that the species of 
Styhphora, to which Dimcan attached the name raristeUa, also inci- 
cates a rather old Tertiary formation. Might these two species have 
been mixed with specimens trom a younger formation ? Having in 
37149— 19— Bull. 108 3 



Digitized by VjOOQIC 



216 BULLETm 108, UNITHD STATES NATIONAL MTT8EUM. 

mind the information above stated, I published the suggestion that 
some of the Santo Domingan fossil corals are perhaps of Pliocene age.' 

Recently Miss Carlotta J. Maury has submitted to me for study the 
fossil corals she collected during an expedition to Santo Domingo. 
She informs me that the zones on Rio Gurabo are lettered in strati- 
graphically descending series, '*A" being at the top and '*G" at the 
base of the section; zone H on Rio Cana is considered to be the same 
as zone G on Rio Gurabo. Bluflf 1 on Cercado de Mao is correlated 
by Miss Maury with a part of the Rio Gurabo section above zone G, 
and bluff 3 on Cercado de Mao is correlated with that part of the Rio 
Gurabo section below zone F. 

As regards the corals, the definite stratigraphic tie-point is foimd 
in zone H on Rio Cana, where three species which also occur in the 
Bowden marl of Jamaica were collected. It has been stated on pp. 
212, 213 of this paper that the Bowden coral fauna is stratigraphically 
above the Oligooene faunas of Antigua, Bainbridge (Georgia), Lares 
(Porto Rico), Empire (Panama), and Tampa (Florida). These Santo 
Domingan corals, except. those from zone G-H, therefore belong 
stratigraphically above the horizon of the Bowden marl. In a manu- 
script now almost ready for press I am describing as new six addi- 
tional species of Placocyathus from Miss Maury's collection. These 
are not entered in the table following. 

I Washington Acad. Sd. Jour., vol. 5, p. 489, 1915. 



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OBOLOOT Ain> FALBONIOLOOT OV THB OAITAL ZOKB. 217 






I 
1 

I 

O 




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218 BULLETIN 103, TJKITED STATES KATIONAL MUSEUM. 

An inspection of the foregoing table shows that at zone H Orbicdla 
cavernosa and Solenastrea bovmonif both now living, were collected, 
bringing the total of living species from the Bowden horizon up to four. 

The following are my conclusions on the geologic age of the coral- 
lif erous beds of Santo Domingo : 

1. The oldest fauna represented by Miss Maury's collection, zone 
H on Rio Cana, is that of the Bowden marl. It is somewhat younger 
than the Chipola marl of Florida and is of BurdigaUan age according 
to European nomenclature. 

2. Zone F is closely related to G and H. It is also probably of 
BurdigaUan age, and corresponds to a part of the Alum Bluff forma- 
tion of Florida lying above the Chipola marl member. 

3. Zone E and D are faunally near the imderlying beds and are 
probably of uppermost Burdigalian or Helvetian age. 

4. Zones C to A, inclusive, are probably of Helvetian age. 

5. The Santo Domingan coral faunas are younger than the exten- 
sively developed Ohgocene coral reefs of (Georgia, Florida, Cuba, 
Porto Rico, Anguilla, Antigua, and Central America. 

6. The presence in Santo Domingo of AsterosmUia exarata variety, 
which is also foimd in the Antigua formation, of a species of Lepto- 
muasa, and of Siderastrea conferta (Duncan) typical, indicates that 
there are deposits of middle and upper Oligocene age in Santo 
Domingo, but Miss Maury did not make collections of corals from 
those horizons. 

CUBA. 

BARACAO AND MATANZA8. 

Fossil corals of Bowden age were collected at two locaUties — 
namely, station 3476, in a yellow marl at Baracao; and station 3461, 
also in a yellow marl in the gorge of Yumuri River', Matanzas. The 
species are as follows: 

Fossil corals from Baracoa and MaUmzaSf Cuba, 

I 



Name. 



Baracoa. MataiiTiw Bowdtn 



St^lophora ffranuUUa Duncaii 

Poduopon bofocodefuii Vau^iaii. 

Madracis mirabttis (Dtichassaing and Mlchelotti). 

Tkfmntu hapeti Vaufban 

PoHUs baroeoHentit Vaudian 



X 
X 



X 
X 



var . matanzasetuu Vaughan . 



LA CRUZ MABL. 

This name is proposed for the bedded, yellow, argillaceous, and 
calcareous marl particularly well exposed on the east side of Santiago 
Harbor between Santiago and the Morro. The type exposures are 
along the railroad eastward from the La Cruz to the crossing of the 
highway from Santiago to the Morro. The corals collected in this 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 



219 



formation are listed below. Descriptions of the moUusca by C. W. 
Cooke will appear in a forthcoming publication of the Carnegie In- 
stitution of Washington. The corals are as follows: 

Fossil corals from the La Cruz marly Cuba. 



Name. 



Stfflopkora a/finU Duncan 

Poduoporo species 

Sttphanoeoenia inUT$ep(a (Esper) , 

OmceUa limbata (Dtmcan) 

SoUnattrea hpades (Dana) 

botimoni M. Edwards and Haime . 

TkjfMnus afl. T. aeentrkut Duncan , 

^iite7o«CrM tJd«rM (Ellis and Solander) 

Gonhpora jacobiana Wax^gjtiasi 

/>orft«portte»( Pallas) 

aatreoide* (hamack) 



.Santo 

Domingo 

above 

tone H. 



X 
X 



Santo 

Domineo 

zone H 



Bowden. 



Recent. 



X 
X 



X 
X 



Of 11 species listed above, 5 are now living in the Antillean re- 
gion; but of the 8 genera represented, 4, i. e. 50 per cent, are now im- 
known in the Atlantic Ocean. The horizon appears to be above that of 
the Bowden marl, and to bo near zones D and E of the table on 
page 217. I obtained niunbers of poor prints and casts of corals 
near or at the base of the formation in the vicinity of Santiago. 
Although they are too poor for determination, they resemble in 
form the species of PldcocyathuSy Asteroarnilia, AntiUia, ThyscmuSf 
and SyzygophyUiay of the Santo Domingan deposits. Similar poor 
casts and imprints suggest that this is a widely distributed formation 
in Cuba. 

FLORIDA. 



ALmC BLtTFF 70RMATI0N. 



The coral fauna of the Chipola marl, member of Alimi Bluff forma- 
tion is small, comprising four species representing as many genera, 
namely, /SfyZopAora, AntUlia, a new genus that resembles a Thysanus 
with a commensal sipunctdid worm in its base, and Ooniopora. 

The coral f aima of the Alum Bluff formation is meager. Exclud- 
ing the Chipola marl member it comprises the foUowing species: 
Fossil corals from the Alum Bluff formation. 



Name. 


Oak 
Grove. 


White 
Springs.! 


Tampa 
brick- 
yard. 


Attrkilia new species 


X 








X 


X 


tiJtcfnH* Vffughan , . 




X 


Oonkroon iacobkina VauKhan , ...... 




X 











< For descriptkni of the stratigraphic relatk>ns of beds at White Springs see Vaughan, T. W., and Cooke, 
0. W., Corrilatton of the Hawthorne fonnatk>n, Washington Aoad. Sci. Joum., vol. 4, pp. 360-253, 1014. 

Although, in my opinion, the formation in which these corals occur 
should be referred to the Miocene, I beUeve it is very low Miocene, 



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220 



BULLBTIN 108, UNITED STATES NATIONAL MTTSEUM. 



bdow the Bowden horiion. A recent discovery by Sellards is of 
importance in determining the age of the Alum Bhiff formation.* 
The following is a list of the vertebrates: 

Pardhippua leonensis Sellards. 

Merychippua species. 

Mesocyont leonensis Sellards. 

OxydactyliLsf 

Leptameryxf 

Sellards says: 

It would seem, therefore, as a whole, that the vertebrate fossils indicate that the 
Alum Bluff formation is to be referred to the Miocene. The presence of prot<^ppine 
horses in particular would seem to be decisive as to the age of the formation, exclud- 
ing its reference to the Oligocene. 

The opinion of Prof. J. C. Merriam on the age indicated by the 
Merychippus is quoted. He says that he would judge the horizon 
to near the lower portion of the middle Miocene. Later Professor 
Merriam informed me that he considers the Merychippus as of lower 
Miocene (BurdigaUan) age. 

The evidence in favor of considering the Alum Bluff as of lower 
Miocene i^e m^ht be greatly multiplied. The presence at Oak 
Grove, Yellow River, Florida, of a species of Astrhdia closely related 
to A. paimata (Groldfuss) of the Maryland Choptank and Calvert 
formations suggests Miocene. Pecten myanus Dall indicates Mio- 
cene. Canu and Bassler are positive that the Bryozoa are of Miocene 
age. Berry's opinion based on his study of the fossil flora ' is not 
incompatible with this interpretation. 

MIDDLE AND SOUTH ATLANTIC STATES. 

The following is a list of the Miocene species, as far as at present 
known:' 

Miocene corals from the Middle and South Atlantic Stales. 









Geologic (ormaUon. 




Name. 


Calvert. 


Ohop- 
taiiK. 


* St 
Marys. 


l^. ': ^^ 


ChocUw- 
hatdiee. 


Pvnewthus vaugkaiU Oane. 




X 


y X 




AstrlieUa wilmata rOoldfaas) 


X 


X 


....f.....i -:---.:- --. 




Aatramtia lineata ^Conrad) 


X 
X 
X 


V i 




eowndi VauiEhui 






^ ; ! 


SeottMrett marvlandiea (Conrad) 






X 


X 


crouM rTiioniAV and TToIrmni) 






X 




liivHa vauahani (Qr^gorv) 




















'1 





i Sellards, £. H., FossU vertebrates from Florida, A new Miooene teuna, Florida GeoL Sorv., Sth Am. 
Hept., pp. 83-92, 1916. 

t Berry, £. W., The physloal conditions and age Indicated by the flora of the Alom Bloff formatioo, 
U. 8, OeoL Survey ProL Pap. 98-E, pp. 41^9, pis. 7-10, 1916. 

* Vanghan, T. W., Anthosoa: ICaryiand Qeol. Survey Miooene, pp. 188-448, pis. ia»-l», 1904; The reef 
coral fauna of Carriso Creek, imperial Oounty, Oalifomia, and its significance, U. 8. OeoL Snrvey FroC 
Pap. 9S-T., p. 366, 1917. 



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GEOLOGY AND-PALBONTOLOGY 07 THB OAITAL ZONB. 221 

Berry has recently reviewed the Miocene Calvert flora of Maryland 
and Virginia; and expressed the following opinion:^ 

Seven of the Calvert plants, or 26.9 per cent, are common to the Tortonian of Europe, 
and 10 others, or 38 per cent, are represented in the Tortonian by very similar forms. 
In view of the fact that these floras spread into both regions from a common and equally 
accessible source, as I have just stated, the evidence that the Calvert flora indicates 
a Tortonian age is as conclusive as intercontinental correlations can ever be. Com- 
pared with other American floras of Miocene age, that of the Calvert has little in com- 
mon with the described Miocene floras from Colorado, Idaho, Oregon, or California, 
which are all lake or river valley floras of moist upland forest types. 

Should Berry be correct in his correlation of the Calvert with the 
European Tortonian,. there is at present no definitely recognized 
Helvetian Miocene in the Coastal Plain of the United States; and con- 
sequently no Helvetian coral-fauna. 

COSTA RICA. 

Corals representing the Bowden horizon or one very near it were 
obtained in Costa Rica at two locaUties, viz: 

'*Liinon, CoUine en dfimoUtion,'' No. 618 of the H. Pittier collection; 
and at station 6249, Hospital Point, Bocas del Toro. The species 
from the former of these locahties are as follows: 

AsterosmUia hiUi Vaughan. 
Stephanocoenia intersepta (Esper). 
Dichocoenia iuberosa Duncan. 
BdlanophyUia pUtieri Vaughan. 

BalanophyUia pittieri was obtained at Hospital Point as well as 
at Port Limon. 

PANAMA. 

The type of Stylophora portobellensis Vaughan, from Portobello, 
was probably collected in the Gatun formation. 

COLOMBIA. 

Mr. George C. Matson collected at a locality 0.5 kilo meter east of 
Usiacuri in association with a fauna representing the Gatun forma- 
tion specimens of Septastfea maisoni Vaughan, which is very nearly 
related to Septastrea marylandica (Conrad)— a species common in 
the St. Marys and Yorktown Miocene of Virginia. The available 
evidence leads to the opinion that the Gatun formation is of Miocene 
age, and that part of it is of upper Miocene age. 

CONCLUDING REMARKS ON THE MIOCENE. 

The Gatun formation, the formation next above the Emperador 
limestone, according to the geologic map, plate 153, occurs only on 
the north flank of the Isthmus and does not extend from ocean to 
ocean. There is in the Canal Zone no evidence to indicate inter- 

» U. 8. Oeol. Survey Prof. Pap. »-P., p. 66, 1916. 



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222 BULLETIN 103, UKITED STATES NATIONAL MUSEUM. 

oceanic connection during Miocene time, although there was such 
connection in other areas not far away, in Nicaragua for instance. 
During the Miocene there was a very weak development of reef- 
corals in Central America, the Antilles, and the southeastern United 
States, as the foregoing lists show. The Miocene is characterized by 
the disappearance of many genera of corals that were abundant in 
the middle and upper Oligocene and by the introduction of the modem 
coral-fauna. However, a number of genera at present known Uving 
only in the Indo-Pacific persisted. These genera are as foUows: 
Placotrochus, PociUopora. SyzygophyUia. 

Placocyathus. AntiUia, Pavona. 

Siylophora. Favites, . Ooniopora. 

Of the Miocene genera, Astrhdia, Septasireaj and Thysanus are not 
known Uving. 

PUOCENE. 
CALOOSAHATCHEE MARL, FLORIDA. 

The following species of corals have been recognized in the Caloosa- 
hatchee marl: 
*Archohelia limoTiensis Vaughan. 
Dichocoenia new species 1. 
new species 2. 
Meandrina maeandriies (JLiinnaeus). 
Cladocora johnaoni 6ane. 
PhyUangiaJhridana Gane. 
*Solena«trea Jiyades (Dana). 

Houmoni M. Edwards and Haime. 
Septastrea crassa (Tuomey and Holmes). 
Thysanus species. 
Maeandra pliocenica (Gane). 

afif. M, strigosa (Dana). 
aff. M. divosa (Ellis and Solander). 
*8idera8trea pliocenica Vaughan. 

*daUi Vaughan. 
*PoriU8 poritea (Pallas). 
"^furcata Lamarck. 
dvoaricata Le Sueur. 
Those species whose names are preceded by an asterisk are con- 
sidered in the descriptive part of this paper. 

The foregoing list is complete for the Caloosahatchee corals from 
Oaloosahatchee River and Shell Creek, Florida, except one species 
of whose genus I am not sure. There are in the United States 
National Museum 19 species from the Caloosahatchee marl. Of 
these 19 species, 6 and perhaps 8 are also Uving in the Floridian 
region, while the other species, except those belonging to Septastrea 



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(»OLOQY AND PALEONTOLOGY OF THE OANAL ZONE. 223 

and ThysanuSf have close relatdves in the present Floridian fauna. 
I have previously pointed out^ that this fauna contains no genera 
at present confined to the Indo-Pacific, such as PlacoirochuSy Placo- 
cyoUhuSy Siylophora, PocxUopora^ AntUliay Syzygophyllia, and Oonio- 
pora, all of which occur in the West Indian Miocene, and all except 
the first two also occur in the West Indian Oligocene or Eocene. 

LIMON, COSTA RICA. 

Certain corals collected in the vicinity of Limon are reputed to 
come froiii a bed of Phocene age. They are as follows: 

Madrads mirabUis (Duchassaing and Michelotti). 

Archohelia limonensis Vaughan. 

OrbiceUa annularis (EUis and Solander) var. 
cavernosa var. endoihecaia (Duncan), 
var. cylindrica (Duncan). 

Except Archohelia limonerhsiSj it appears that these corals might 
represent the Santo Domingan Miocene above the Bowden horizon. 
The material is not adequate for a positive opinion. 

CARRIZO CREEK, CALIFORNIA. 

Recently I have described in detail an interesting small reof- 
coral fauna from Carrizo Creek, Imperial County, California.^ 

The following table, taken from the paper mentioned, contains 
the names of the species composing this fauna and of the most 
nearly related species in Florida and the West Indies. 

Coraitfrcm Ccarizo Creeks Col. 

fj^fff^ Most nearly related species in Florida or West 

^^~"* I Indies. 

AiMiUia carrixauit Vaoghan I JStwmttia/iuei^toto (Pallas), PI, R. 

Didtoeoenia merriami (y^g^ian) Wkkocoenia specira, P; D. stokesi Milne Edwards 

var. croMisepto Vau^ian {/ and Haime, PI, R. 

Solenastrea fairbankti (Vaughan). typical | 

JS*2l!t!ISyvi.^Jh5?*"^ \\stUMUtrea h^aiet (Dana) and 8. hawwmi Milne 

:Si-SZr^^JgaS?^ • }Edward>a,;7HSn;e,P,HR. 

JToaaiMlra bow«r«i Vaughan I Maeandra Utbyrinthiformia (Linnaeus), PI, R. 

^'^'TSPSS^SfWiS'^::::::::::"::^ v.u«imn, p. 

Siiemtrm etiifwnka\augjaan SUeroitrM pUoeenica Vaughan, P . 

Porttit c«rr<2»uif Vaughan. " . -- - - ~. 



Porites attreoidet Lamarck, PI, K. 



P, Pttooone; PI, Pleistocene; R, Recent. 

Regarding the geologic age of this fauna, it was said: 

The specific affinities of the Carrizo Creek corals are discussed in detail after the 
descriptions in the systematic part of this paper. The Carrizo Creek species are so 
near species belonging to the same genera in the Pliocene Caloosahatchee marl of 
Florida and in the Pleistocene and living reefs of Florida and West Indies that it 
seems to me they can scarcely be so old as Miocene; lower Pliocene appears to be the 
maximnm age which may be assigned to the fauna. 

1 The reelHSoral teuna of Carriio Creak, Imperial County, Califoniia, Prof. Pap. OS-T. p. 3S0, 1017. 
« U. 8. Geol. Surrey Prof. Pap. »-T, pp. 356-38S, pl». W-102, 1917, 



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224 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

The following is said as to the bearing of this fauna on a possible 
post-Oligocene interoceanic connection: 

That there was interoceanic connection across parts of Central America daring 
upper Oligocene time and' that this connection was terminated in Miocene time is 
geno^y admitted. The extinction of Pacific launal elements in the Gulf of Mexico, 
the Caribbean Sea, and the Western Atlantic Ocean has been discussed and sum- 
marized on page 366. Was there interoceanic connection during upper Miocene or 
Pliocene time after the sharp differentiation of the Caribbean and Mexican Gulf 
faunas from the Indo-Pacific faunas, thereby permitting interoceanic faunal migra- 
tion? The discovery of a reef-coral fauna of piu^y Floridian and Caribbean faciee at 
the head of the Gulf of California strongly suggests, if it is not positive proof, that the 
western Atlantic fauna extended from the Atlantic into the Pacific after the faunal 
differentiation had taken place. It is well known that the living reef -coral ^una on 
the Pacific side of Central America is depauperate in comparison with that on the 
Atlantic side. Greater vigor may account for the dominance of the migrant fauna 
over the Pacific fauna, which was finally suppressed, or geologic or other ecologic 
conditions that are not yet understood may have excluded the Pacific fauna from the 
head of the Gulf of California, while they permitted the migration of the Atlantic 
fauna into that area. 

That the suggested interoceanic connection existed can scarcely be doubted. To 
locate it, in the present state of meager knowledge of the areal and stratigraphic geology 
of Central America, is not possible. Perhaps it was across the Isthmus of Tehuante- 
pec. The problem awaits future investigation. 

This fauna diflFers from the Miocene fauna of the La Cruz marl of 
Cuba in the absence of genera at present living in the Indo-Pacific, 
for instance, StylopTwra^ PocUhporaj and Gonioj>ora. As none of the 
Indo-Pacific genera occurs in the Carrizo Creek fauna, and as only- 
genera of Atlantic affinities have been foimd there, it seems neces- 
sary to infer that the fauna migrated from the Atlantic to the head 
of the Gulf of California after the Indo-Pacific genera had become 
extinct in the Atlantic. This would mean connection between the 
Atlantic and the Gulf of California in very late Miocene or Pliocene 
time. 

Attention should here be called to a statement for which I am 
responsible. It is said in the report referred to below * that some 
fossils obtained by Mr. William Palmer in a quarry in Calle Infanta, 
Habana, may be of Pliocene age, although it is probable that they 
are Pleistocene and that other limestone near Habana is perhaps of 
Pliocene age. The material obtained by Mr. Palmer is very poor, 
but some specimens are casts of the inside of the calice and the inter- 
septal loculi of a large bilobate species of AntiUia. The species more 
probably is A. walli Duncan of the Bowden marl, but it might be 
A. hUohata Duncan; another cast seems to represent a species of 
Thysanus; while another is a species of SyzygophyUiay probably 
S. dentata (Duncan). One specimen of Stephanocoenia intersepta 
(ElHs and Solander) is identifiable. The material seems quite 
clearly to represent either the Bowden or a somewhat higher horizon 

» Hayes, C. W., Vaughan, T. W., and Spencer, A. C, A geological reconnaissance of Cuba, p. 23, 1902. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 226 

in the Miocene. It i&not Pliocene, according to our present knowledge 
of Pliocene coral faunas. 

Pleistocene. 

Only the names of the Pleistocene species considered in this paper 
are given in the following lists: 

Pleistocene corals from Mount Hope and Colon y Canal Zone. 

OcvUina diffusa Lamarck. 

varicosa Le Sueur. 
EusmUia fasiigiata (Pallas). 

Asirangia (PhyUangia) amerieana M. Edwards and Haime.* 
Cladocora arbuscida Le Sueur. 
Solenastrea boumoni Wine Edwards and Haime. 
Faviafmgum (Esper). 
M<ieandra areolata (Linnaeus). 
Manicina gyrosa (Ellis and Solander). 
Agaricia agariciks (Linnaeus). 

var. purpurea Le Sueur. 
pusUla Verrill. 
Sidera^trea radians (Pallas). 

siderea (Ellis and Solander). 
Acropora muricata (Linnaeus)* 

palmata (Lamarck) at Colon. 
Porites furcata Lamarck. 

oMreoidfs Lamarck. 
MiUepora aldcomis Linnaeus. 

It will be remarked in passing that the coral fauna at Mount 
Hope is a typical inner-flat coral fauna. 

PJeistocene specimens were obtained at Monkey Point and limon, 
Costa Rica. The list is as follows: 

Pleistocene corals from Monkey Point and Limon, Costa Rica, 



Name. 



EvsmiUa/attiffiataCPaXiaa) 

M^eanif aivota (Ellis and Solandar) . . 

8Mgo$a(ptaiA) 

Mknieina §ifTO*a (Ems and Solander)... 
A^rMa ugarkitet var. era$ta Verrill . .. 
SiierttTM 9iieff (ElUs and Solander) . 



Monkey 
Point. 



X 
X 
X 



X 

Aeropors murkaiaCUjxDaeoa) X 

poimoto Lamarck X 

Pori<M /urccfa Lamarck t 



Limon 
MQinHiU 



X 
X 



The corals from Monkey Point represent a seaward-facing reef; 
while those from Moin Hill are more characteristic of inner-flat 
conditions. 

1 Names added in Uie proof and not entered in the table o( spedesi pp. 228-287., or the systematic 
aoooont of the faunas. 



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226 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

SUMMABT OF THB 8TBATI0RAPHIC AND QBOQRAPHIO DISTRIBUTION OP THB TSBTIART 
AND PlBISTOCENB CORAL-FAUNAS OF CENTRAL AMERICA AND THE WeST InDIES. 

1. The upper Eocene coral-fauna of the St. Bartholomew lime- 
stone is known in St. Bartholomew, in Jamaica, and on the Pacific 
side of Nicaragua. 

2. No lower OUgocene coral-fauna is at present known in the 
West Indies or Central America. 

3. Rich middle OUgocene coral-faunas are known in Antigua, 
Porto Rico, Cuba, Georgia (near Bainbridge), Alabama (Salt Moun- 
tain), eastern Mexico, Panama, and the Island of Arube. The same , 
fauna is known to be present in Santo Domingo. 

4. Upper OUgocene coral-faunas are present in Anguilla, the 
Canal Zone, Florida (Tampa formation), and there are some reef- 
corals representing the same fauna in Cuba. There seems to be a 
distinct break between this and the succeeding Miocene faunas. 

5. The Bowden, Jamaica, lower Miocene fauna is represented in 
Santo Domingo, Cuba, and Costa Rica. This fauna is probably 
yoimger than the coral-fauna of the Alum Bluff formation in Florida. 

6. A closely related but highw Miocene fauna is present in Santo 
Domingo and Cuba. It seems probable that this fauna is geologi- 
cally older than the coral fauna of the Maryland and Virginia Miocene. 

7. The presence at Usiacuri, Colombia, of a species of Septastrea, 
very closely related to S, marylandica of the St. Marys and Yorktown 
Miocene of Virginia, suggests the presence in northern South America 
of a middle or an upper Miocene coral fauna. 

8. There is a moderately rich PUocene faima in the Caloosahatchee 
marl of Florida, and this faima appears to be represented at Limpn, 
Costa Rica. 

9. Pleistocene reefs are extensively developed in Central America, 
the West Indies, and Florida. 

10. Living reefs exist in the same areas in which there are Pleis- 
tocene reefs. 

11. The periods of reef-coral development are as foUows: 

(a) Upper Eocene St. Bartholomew Umestone, weak development. 
(6) Middle OUgocene, the greatest known development of American 
coral-reefs. 

(c) Upper OUgocene, considerable development of reefs. 

(d) Miocene, weak development of reefs. 

(e) PUocene, weak development of reef-corals in Florida. 
(/) Pleistocene, extensive development of reefs. 

(g) Recent, extensive development of reefs. 

12. Periods of connection between the Atlantic and Pacific oceans 
are as follows: 

(a) Upper Eocene. 

(b) Middle and upper OUgocene and lower Miocene. 

(c) A connection, probably narrow, in very late Miocene or in 
PUocene time. 



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228 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Table op Stsatigraphic and 



Name of species. 



Eocene— Brito 
formation, 
Nicaragua. 



OUffooene. 



Horison of 

Antigua 

formauon. 



Culebra 
formation, 
Canal Zone. 



Horizon of 

AngoUla 

formation. 



Stplophora HnperatorU Vaughan.. 



tfd Cut; AnguUla. 
mL 8. I 



panomtntU Vaughan. 
a^nif Duncan 



.1 aaOlard Cut; 
U mL 8. 
Monte Lirio. 



portobeUentiiVBUgban. 

goethalti Vaughan 

macdonatdi Vaughan.. . 
granuUtta Duncan 



canalit Vauxhan — 
po4i4«roM vaughan . 



PodUopon amoldi Vaughan ^ . . 

&or<M»^i«n#i« Vaughan 

gtMntanameniit vaughan. . 

liairaeit mtrabilU (Duchaasaing and 

Hichelotti). 
Attroeoenia d*aekiardii Duncan.' 



ffuantanamentii Vaughan . 



Also St. Bar- 
tholomew. 



incruttafu (Duncan). . 
dectttureruU Vaughan . 



meimeri Vaughan 

porUnicensis Vaughan.. 
Stifloeoenia pumpeU^i (Vaughan).. 
Ocuiifw diffusa Lamarck 



oaHcoM Le Sueur 

ArehoMia Ifmonentii Vaughan . 



AttmtmmakmiVhughui 

StephMocoenia intenepU (Esper) . 



DidiceoenU Mema Donoan. 



BuimUia/iutlffiQia (Pallas).. 



Cladocora arfrutevte (Le Sueur) 

OrMeeOm afmularis (Ellis and Sol- 
ander). 
Umteto (Duncan) 



J U mi. S. of 
Monte Lirio. 



Alabama; An- 
tigua. 



Near Ouanta- 
namo, Cuba. 



Antigua; near 

Quanta namo, 

Cuba; Tono- 

sl, Panama. 



Bainbridge, 
Oa.; Antigua; 
nearOuanta- 
namo.Cuba. 

Near Quanta- 
namo,Cuba. 

Anttam; Lares 

Bain bridge, 
Ga.; Antigua. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 229 

GsoGRAPHio DiaTBiBunoN OF Species. 



oUgomDe— ' 
Continued. 



Empendor 
limesloDe, 
CaaftlZone. 



I 



ICiooene— 

Horisonof 

Bowden 

marl, etc. 



Pliocene. 



Pleistocene. 



Recent. 



Remarks. 



Empire; Rio 
AfnaSdhid. 

Empire 



Santo Domin- 
go; Santiago, 



Empire 

Empire; Rio 
AguaSahid. 



Empire. 



Matansasand 
Baracoa, 
Caba; Rio 
Qurabo, St. 
Domingo,etc. 



Empire. 



Rio AguaSa- 
hid. 



Baracoa^Cuba 



Matansas, 
Cuba. 



Limon, C. R.; 
Boivden, Ja- 
maica. 

Bowden, Ja- 
maica; Ll- 
mon, C. Re 
Santo Domini 
eo; Santiago, 

Limon, C. R.; 
Santo Domin- 
go. 



Santo Domin- 
go, Cuba. 



Colon, Costa 
Rica. 



C. R.; 
Calooia- 
hatoheemarl, 
Florida. 



MtHope,C.Z. 
Mt.Hope,C.Z. 



West Indian 
region. 



Limon,C.R... 



Mt. Hope. C. 
Z.; Monkey 
Pt., C. R.; 
W.L 

lCt.Hope,aZ. 

W.L; Fk.; etc. 



Porto Bello; prob- 
ably Gatun forma- 
tion. . 



Caribbean re- 
gion. 



Upper Eooene of St. 
Bartholomew. 



Florida; West 

Indies, etc. 

....do.. 



West Indies, 
Bermnda,eto. 



West Indies; 
Florida, etc. 



Very near the Plei- 
itooeae and living 

D, 9toke»i M. Ed- 
wards and H<dfi i> 
of the West Indies, 
Florida, etc. 



....do 

....do 



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230 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Table op Stbatiora pmc and Gbooraphk 



Name of species. 



Orbicella imperatorU Vaughao.. 

antiUarum (Duncan). 
aUi99ima (Duncan) . . . 



cavemoia (Linnaeus) 

var. endothecata (Duncan) . 

var. cvlindriea (Duncan)... 

aptfrto (Verrill) 

bainbridgeruis Vaughan 

cottata (Duncan) 

canolif Vaughan. 



var. tiUcentis Vaughan . 
brevU (Duncan) 



ifuiffnit (Duncan) 

intermedia (Duncan)., 
ffobbi Vaughan 



imdiani (M. 
Halme). 



Edwards and 



Solenastrea hpadet (Dana).. 



boumoni M. Edwards and 
Haime. 



Septattrea matsoni Vaughan 

AntiguaUrea cellulota (Duncan).. 



var. curvaia (Dunoan). , 
var. aileeeruis Vaughan. 



elegans (Reusa) 

alveolaris (Catnllo).. 

Stjflanifia panameruit Vaughan . . 
Faviafragum (Esper) 



nuiedondldi Vaughan . . . 

Fmitet madcaiM Vaughan .. . . 

polpgonalit (Duncan).. 



OonUutrea canalis Vaughan 

Maeandra antiguentii v aughan . 



Eocene— Bri to 
formation, 
Nicaragua. 



portoricen»is Vaughan. 



dumblei Vaughan . . . 
areolata (Linnaeus) . 



OUgocene. 



Horixonof 

Antigua 

formation. 



Bain bridge, 
Qa. 
Antigua; Lares, 



Culebra 
formation, 
Canal Zone. 



Hofiionof 
Angidlla 



Angullls, 
Cuba. 



Antigua; Am- 
be. 
Antigua 



Antigua; P. 

R.; Cuba; 

Oa.; eastern 

Hex.; Arufoe. 

Antlgoa 

Bain bridge, 

Qa.; Antigiia. 



Tonosi, Pana- 
ma; Antigua. 

Eastern Mexi- 
co. 

Antigua; Bain- 
bridge, Oa. 



Antigua; To- 
nosi,Panama. 

Lores, Porto 
Rico. 

Eastern Mez.. 



LasCasoadas. 



Anguflla. 

Angnflla. 



Anguills 
Tampa, Flori- 
da. 



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GBOLOGY AND PAUBOKTOLOOY OF THE OAKAL ZOKB. 231 

Distribution op Spboiks— Continued. 



OUgooene— 
Continued. 


Miooene— 

Horltonof 

Bowden 

marl, etc. 


Pliocene. 








Smpoiadof 
limestone, 
Canal Zone. 


Pleistocene. 


Recent. 


Remarks. 


Empire and 
near Ifira- 
flores. 




1 




. 






Montsarrat, geologic 










horixon unknown. 
St. CroU, Trinidad; 









West Indies; 
Florida. 


West Indies; 
Florida; Bra- 
sil. 


probably about the 
horixon of the An- 
tigua formation. 


I 


Santo Domin- 
go. 
,do • .. 


T4mon,C, R. .. 






....do i 




!.. . >...... ... 




:::::?!^..... !.::!::::..;.:::..::. Brasu 






go. 






i 


.1 1 




Emoire 




i ; 










1 


Tampa, Fla.; about 


1 









the horison of the 
Anguilla formation. 
Do. 


1 








Santo Domingo, Niv- 
i^ shJeTHoriion 
unknown. 








i 

t 
t 



















;!*.;!.*;;;;;;.;.■ 


Santo Domingo; hori- 








1 

1 


lonunknoiwh. 
Lutetian (Eocene); 


1 


Santiago and 
Citfnaga, 
Cuba. 

Santiago, 
Cuba: Santo 
DomiJDgo. 

Usiaciirf, Co- 
lombto. 


Caloohat- 
cjb^ marl, 

do 


Wett Indies; 
Fl(Hflda. 


West Indies; 
Florida. 

do 


Rupelian ' (Olioco- 
oene) of Veneto, 
Italy. 














Formation in part the 










sfune as the' Oatun 
formation. C.Z. 
Byram marl, Miss. 






































1 








*" 

Empire, C. Z. . 














-^^rȣ'= 


W. I.; Fla.; 
Bermudas; 
Asores; St 
Vincent. 




























Empire, C. Z. . 














;/;* .. 






























Mt.Hope,C.Z.; 


W. L: Fla.: 


i 






W. ifiFla., etc. 
etc.- ' 



37149— 10— Bull. 108 i 



Digitized by VjOOQIC 



232 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

TaBLB or STBAnORAFHIO AND GSOORAFHIC 



Nameofspeoiee. 



Mamnira eUvota (Ellis and SoUndflr) . 

ttriffota {Dnna) 

LeptortatpeneeriYKOghan 

Manidna gprom (Ellis and Solandor) . 



tBittouckUentiiywigjban. 
Tkptanut excentriau Duncan 



hapetiVaagltian.. 
r meifueri Vau^ian. 
Aforicia agturMtet (Linnaeus). 



fimjIMpftfOto hoiftti Vau^ian. 



Tar. purjntrea Le Sueur. . 
Tar. eroMs Verrill - 



TBT.piMlUaVerriU... 
anffuUtentU Vaoghan . . 
iominkentii Vanghan. . 



PoooMjwnaiiwMfoVaiichan 

ZrCptoMTif porftwieeiuit vaogban . . . 
Pinnattraea unfuiUenrit Vaugban , 

anUguentii Vaugban. . 

Siierulrea portoiie (Duncan) 



I (Pallas).... 

KeOstoVeniU 

cpi^AfM (Duncan).. 



p(mrtaUH Vaugban 

pliooentoa Vaugban 

JUOfftoroeiwif Vaugban. . 



tidern (Ellis and Solander) 



Tar. diminkefuit Vaugban 
tiUeentU VAUghan 



do/U Vaugban. 



Eocene— Brito 
fDrmatkm, 
Nicaragua. 



Brito. 



OUgocene. 



Horlsonof 

Antigua 

formaiion. 



Culebra 
fSormation, 
OanalZone. 



R to Canapu, 
c;uba; Anti- 
gua?. 



Antigua.. 



Quantana m o; 
Tonoai, Pana- 



Lares, P.R... 
Lena road, 

ZoneC,P.R. 
Antigua; (hian- 

tanamo,Cuba. 



Horiionof 
Angidlla 
formation. 



Anguilla. 



Angunia. 



Anguilla. 



Digitized by VjOOQIC 



QSOLOGY AND PALEONTOLOGY OF THB OANAL ZONE* 233 

DisTBiBUTiON OF SPBCiBS — Oontiiiued. 



ConUnued. 


Miooene- 

HorUon of 

Bowden 

marl, etc. 




■ 


Recent. 
do 




Emperador 
limestone, 
Canal Zone. 


Pliocene. 


PWstocene. 


Remarks. 









Monkey Pt., 
C. B.; W. L; 
Fla. 

do 










do 






















Mt.H(»e.CJS.; 
Mdnmll.C. 
E.;W.L;etc. 


Fla.; W.I.; etc. 














Santiago, 
Cuba: Bow- 
den. Jamaica. 

Matanias, 
Cuba. 
















































etc 
MtHope.CZ.. 


W. I.; Fla.; etc 
, ...do 


















Moin HiU, 
C.B. 

Mt.Hope,C.Z.. 


Especially 
abundant 
east side of 
Andros Is., 

Colon, C.Z.... 






















Santo Domin- 










Emnlre 
































. 
























St. Croix,^Trlnidad; 
probably nearly the 
same as the Antigua 
horison. 








etc 


W. L; Fla.; 
Bermudas. 

Braiillan 
reefs. 


















Do. 












Santo Domingo; hod 
Km unknown. 

S 

Chattahoochee, Tam- 






Caloosahatchee 
marl, Fla. 

















Bowden, Ja- 
maica; Santo 
Domingo; 
Santiago, 
Cuba. 




Mt.Hope,C.Z.; 
Fla. 


W. L; Fla.; 
etc. 

Haiti 


pa, and Alum Bluff 
formations, Florida 
and Georgia. 



















Do. 






Calooeahatdiee 
marl, Fla. 



















Digitized by VjOOQIC 



234 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Tablb of Stratiobaphic and Geograpbic: 



Name of species. 



Sidenutrea eonferta (Duncan) 

Cifathomorpha rochettiita (Mlchelin) . 



JUWVaujshaa 

&roiim< Vaughan 

fteUtVaughan 

splendent Vau^lan 

angtUUeneie Vaughan.. 
roxborougki Vaughan.. 
anHgueneii (Duncan).. 



(«ntii« (Duncan).. 



Diptoaetrea heliopora (Lamarck). 



craeeolamellata (Duncan).. 



var. magnifiea (Duncan). 



var. nugenti (Duncan) . 
*B]anopkyUia piuieri Vaughan 



Aeropora panameneu Vaughan.. 
ealmdenti* Vaughan . . . 



mtiricata (Linnieus). . 



patmata (Lamarck). 



Attreopora goetkaUi Vaughan 

anUfftteneit Vaughan. . 



portofieentieV, 
Actinaeie tMbamieneie Vaugi 



(/onjopora kitU Vaughan 

panamen^t Vaughan. . 
decaUwentU Vaughan . 



regviarie (Duncan). 



var. micToeco^ka (Duncan) 
ui Vaughan 



joeoMana Vaugh 



impentoH* Vaughan . . 

cafMlit Vaughan 

portoriceneU Vaughan. 



Eooen^—Brito 
formation, 
Nicaragua. 



OUgocene. 



HoriEon of 

AnUgua 

fonnation. 



Antigua; 
Lares, P. R. 



Antl 



r:; 



.do., 
.do- 



Antigua; 

Lares, P. R.; 

Ouantanamo, 

Cuba; eastern 

Mexico. 
Antigua; 

Lares, P. R.; 

Rio Canapu 

and Guanta- 



Antigua; 
Lares, F. R. 
(Zone C); 
Cuba; Bain- 
bridge, Oa.; 
Tonosi, Pana- 
ma. 

Antigua 
Lares, P. R.; 
Ouantanamo, 
Cuba; Bain- 
bridge, Ga. 

Antigua 



Antigua.. 
....do.... 



Antigua; 
Bainbridge. 
Qa. 

Lares, P. R... 

Ala.: Bain- 
bridge, Oa.; 
Antigua. 



Bainbridge, 
(}a.;Guanta- 
namo, Cuba. 

Antigua; P. 
R.; Anibe. 

Antigua. 



Lares, P. R.; 
Antigua. 



Culebra 
formation, 
Canal Zone. 



LasCascadas. 



Horizon of 
Anguilla 
formation. 



Anguilla. 



Anguilla. 
do.... 



U mi. S. of 
Monte Lirio. 



Anguilla. 



Angoilla. 
do.... 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 285 

Di»TiuBX7TiON OF SPBOiBS—Continued. 



1 

1 oiigoowie— 
Cootinoed. 

. Emperador 

1 limestone. 

Canal Zone. 


Miocene- 

Horixonof 

Bowden 

marl, ete. 


Pliocene. 


Pleistocene. 


Recent. 


Remarks. 


i 












t 










Lattorflan and Rcpe- 
lian (Oligocene) of 
Veneto, Italy. 


1 


























































, 








................ 





























1 








East coast of 
Africa to Fiji 
Islands. 




1 


















! 

1 


. 










1 


Limon and 










1 

1 Rio Agua Sa- 
1 lud,C.Z. 
Rio Agna Sa- 
nd and Em- 
pire, C.Z. 
























Monkey Pt, 
and Moin 
HIU, C. R.; 
W. I.; Pla.; 
etc. 

Monkey Pt., 
C. R.; Colon 
C.Z. 


W.I.; Fla.;etc. 
W.I.;Fla.;etc. 




; 








1 Emnire 
































1 










Empire 












..!7^:.:::::: 



















































S a n t lag o ; 
CHiba: White 
Springs, Fla. 










Emnire. . . . 










.... do. 




* 










1 








• 













Digitized by VjOOQIC 



236 BULLETIN 108^ imiTBD STATES NATIONAL MXTBEITM. 

Table of Stbatiobafhio and GsooRAfmc 



Name of species. 



Goniopora eUveiVmigJban 

aueadentit Vaughan . 
Porite9 portUt (POlas) 



/ttfeato Lamarck 

teracod«fW<< Vaughan . 



Eocene— Bilto 
formation, 
Nicaragua. 



var. mataiuaseiuii Vau^^ian. 

iiofMiJOd Vaughan 

touW Vaughan 

attmUet Lumaick 



panamentit Vaoghan 

anguUUiulM Vaughan 

(Spnarxa) howei Vaughan. 

Tnaedonaldi Vaughan 

Millepota aldcornU Linnaeus 



OUgooene. 



HorisoDof 

Antiffua 

formation. 



Ani 



IT:: 



Cnlebra 
formation, 
Canal Zone. 



Horiaonof 
AngniOa 
formatiaiL 



do.. 



Anguilia. 



AnguUla. 



Digitized by VjOOQIC 



GEOLOGY AND PALEONTOLOGY OF THE OANAL ZONE. 
DifiTRiBunoN OF Spbciss— Continued. 



237 



OUgoeene- 
Continned. 


• 

Horisonof 
Bowden 
marl, etc. 


' PUocene. 


Fleistooene. 


Recent. 




Empendor 
limestone, 
Canal Zooe. 


Remarks. 


Empire 


























Santiago, 
Cuba. 






W. I 


W.I.; Fla.; etc 
do.... 








Mt. Hope, C. 
Z.;lIdnHUl, 






Baraooa,Caba; 
Bowden, Ja- 

lCatanza8,Cuba 








1 






Empire 


- - - - .- 






.....do 










Santiago, 
Cnha. 




Fla, 


'w/'LVFiaVr 
Bermudas; 
BrasU,etc. 


• 


Emolre 






do 










do 


*\ 




.•....■•..••..•«• 




do 


..••••••••••••••1 •••....••. 










1 


Mt.Hope,C.Z. 


W. I.; Fla.; etc. 






1 





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CONDITIONS UNDER WHICH THE WEST INDIAN, CENTRAL AMERICAN, 
AND FLORIDIAN CORAL REEF& HAVE FORMED, AND THEIR BEAR- 
ING ON THEORIES OF CORAL-REEF FORMATION. 

A brief review of the results obtained from a study of American 
Tertiary and post-Tertiary corals in their relation to the larger prob- 
lem of coral-reef formation in general will now be given. In a paper 
recently published * I stated that in my opinion coral reefs should 
be studied from at least the following standpoints: 

1. The corals themselves, to ascertain the ecolpgic conditions under which they 
live or lived, and to distinguish the calcium carbonate secreted by corals from that 
contributed through other agencies. 

2. A complex of geologic processes operafing in the area must be studied, analyzed, 
and evaluated — among these are the agencies other than corals whereby calcium 
carbonate may be taken from the sea water, the probability of the solvent acti<Hi of 
sea water on calcium carbonate, the effects of winds, currents, and waves in build- 
ing, shaping, and destroying banks, and in submarine planation. 

3. The strat3graphic and structural geology of the area, including a careful study 
of the origin of the sedimentary rocks with which corals are associated. 

4. The physiography, espedaUy that of the shore line, that of the land area adjsr 
cent to the shore, and that of the sea bottom from the shore to abyssal depths. 

In the subsequent discussion, after defining coral reef, brief atten- 
tion wiD be given to the following topics: (1) The general ecology 
of reef -forming corals; (2) the more striking hypotheses of the for- 
mation of coral-reefs; (3) the conditions under which the American 
Tertiary and Pleistocene reefs have formed and their importance 
as constructional geologic agents; (4) the conditions under which 
the living reefs of the same area formed and their importance as 
constructional agents; (5) coral reefs of the Pacific Ocean and com- 
parison of them with the American fossil and living reefs; (6) sum- 
mary of condiisions. 

It is needless to say that, as an elaborate discussion of the subjects 
mentioned would require a large volume, it is possible in the present 
connection to give only summary statements. 

DsFiNrnoN op the Term ''Coral Rebp." 

As definitions are essential in this ad in other discussions, the 
expression ''coral reef' will be defined as follows: 

A coral reef is a ridge or mound of limestone, the upper surface of which lies, or 
lay at the time of its formation, near the level of the sea,, and is predominantly com- 
posed of calcium carbonate secreted by organisms, of which the most important are 
corals.' 

1 8om« ahoal-wBter corals from Murray Island (Australia), Cocos-Keeling Islands, and Fanning Island, 
Carnegie Inst. Washington Pub. 213, p. 54, 1918. 

t Vaughan, T. W., Physical conditions under ^diiob Palaoioic coral reefs were formed, BuD. Geol. Soe. 
America, yol. 22, p. 238, 1911. 

238 



Digitized by VjOOQIC 



OBOLOGY AND PALBONTOLOGY OF TH£ OANAL ZONB. 239 

Frequently it is difficult to decide whether or no to apply the desig- 
nation ''coral reef" to richly coralliferous deposits that are obviously 
bedded. However, I am applying it wherever corals of reef facies 
seem sufficiently abundant to have formed appreciable rugosities 
on the sea bottom, although the deposits are bedded. 

Reefs predominatly formed by calcareous algae should be desig- 
nated ''nulUpore" or ''Lithothamnium reefs." However, where 
the proportion of these organisms to corals is so nearly the same that 
,only exact computation will decide between the two, such a reef 
may be designated ''Coral." The expression "reef coral" will be 
applied to corals of the facies usual in reefs; and "coralliferous 
limestone" or "coralliferous beds" will be appUed where corals 
are present, although they may be rare. Rock predominantly com- 
posed of the shells of moUusks, of the tests of foraminifera and 
Bryozoa, and of chemically precipitated calciimi carbonate are ex- 
cluded from the category "coral reefs." 

The restricted use of the term "coral reef" in this paper wiU prob- 
ably be disapproved by a considerable nimiber of investigators,but 
in my opinion it is essential to clear thinking. Limestones are 
initially formed by one of two processes, namely, (1) through chem- 
ical precipitation either by inorganic or organic agencies that lead 
to supersaturation of water with reference to calcium carbonate 
(CaCOt), (2) through the activity of oi^ganisms that cause the pre- 
cipitation of calcium carbonate (CaCO,) in contact with their soft 
tissues. Corals belong to a group of organisms that secrete calcium 
carbonate (CaCO,), that is, cause the precipitation of calcium car- 
bonate (CaCO,) in contact with their soft tissues. Every kind of 
shoal-water calcium-carbonate deposit has been called "coral rock"; 
the moUuscan-shell sands of the Bermudas, the chemically precip- 
Fitated calcium carbonate of the ooUtes of Florida and the Bahamas, 
and limestones composed of the remains of Foraminifera and Bryozoa. 
The terms coral sand and coral mud have been applied to bottom- 
deposits in which there is no coral. To apply the term "coral rock" 
or "coral-reef rock" to all the kinds of limestones indicated would at 
the present time, in my opinion, be willful mental obfuscation. 
The study of the origin of limeetones and the classification of lime- 
stones according to the source of their ingredients constitute a 
scientific problem of great geologic importance, and I believe it a 
scientific duty to break away from a usage that in mast instances 
concealed scientific fact. 

The importance of the distinction between "reef" and the mate- 
rial lying between a "reef" and the shore is particularly discussed 
on page 249. 



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240 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Ecology op Rbef-Formino Corals. 

This subject has received the attention of very many investigators, 
and most of the broad principles have long been known. Darwin 
clearly recognized the difference in growth-form of exposed-reef 
corals and the corals that grow in the lagoons.* This subject has 
been discussed at great length by subsequent investigators, of whom 
I am one, but although facts have been presented in a more or less 
statistical way, the principle of adaptation of growth form to envi- 
ronment was as clearly perceived by Darwin ^as it is by anyone to- 
day. Dana's conclusions on the relations of corals to the tempera- 
ture of the ocean have been modified in only a subordinate way. It 
is scarcely known who first recognized the polymorphism of species 
of corals according to difference in habit. The recognition of such 
vegetative adaptation was at least foreshadowed by Klunzinger and 
Pourtalfts. Brook clearly recognized the principle, and during more 
recent years it has been elaborately discussed by (Jardiner, Von 
Marenzeller, Wood Jones, and many others, including myself. The 
literature on coral ecology is enormous, and probably the ecologic 
relations of no other group of marine organisms are so well known. 

Recently I have published two summaries on the physical condi- 
tions under which coral reefs form,' and have discussed in detail the 
temperature relations of coral reefs in a paper entitled Temperature 
of the Florida Coral-reef Tract.* Dr. A. G. Mayer has given im- 
portant information on some of the subjects of coral ecology in a 
paper entitled Ecology of the Murray Island coral reef;* and I 
have given considerable data on the relation between the growth- 
form of colonies and habitat in my monograph, Some shoal-water 
corals from Murray Island (Australia), Cocos-Keeling Islands, and 
Fanning Islands.' The la?t-mentioned paper contains a complete 
bibliography of my publications on corals and coral reefs up to 
March, 1917. 

In the second of my papers referred to in the preceding paragraph,' 
I state on page 99: 

The conditions necessary for vigorous coral-reef development may be summanzed 
as follows: (1) Depth of water, maximum, about 45 meters; (2) bottom firm or rocky, 
without silty deposits; (3) water circulating, at times strongly agitated; (4) an abun- 
dant supply of small animal plankton; (5) strong li^t; (6) temperature, annual mini- 
mum not below 18° C; (7) salinity between about 27 and 38 parts per thousand. 

To this should be added the statement that the mean temperature 
of the coldest month must not be lower than about 21^ C. 

1 Stnictuie and distribution of ooral reo£s, ed. 3, pp. 1-19, 1889. 

• Vaoj^ttn, T. W., Phjrsloal ooodittons under which Paleosoic oonl reeb were fonned, G«oL Soo. Amer, 
Bull., YoL 22, pp. 238-252, 1911; The results of investigations of the eoology of the Floridlan and Bahaman 
shoal-water oorals, Nat. Acad. Sot Proc., vol. 2, pp. 95-100, 1916. See also Corals and the formation of 
ooral reef^, Smithsonian Ann. Kept for 1917 (in press). 

• Oamegie Inst. Washington Pub. 213, pp. 321-339, 1918. 
« Idem, pp. 1-48, pis. 1-19, 1918. 

• Idem, pp. 49-233, pis. 20-98, 1918^ 



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OBOLOGY AKD PALBOKTOLOGT OF THE OANAL ZONB. 241 

Wherever there are well-developed fossil coral reefs it seems safe 
to infer that the physical conditions above enumerated prevailed. 
It is imhecessary to discuss separately each item entered under No. 
11 of the summary statement of the periods of coral reef develop- 
ment on page 226. During upper Oligocene time (the time of the 
deposition of the upper part of the Chattahoochee formation) tropical 
conditions extended in Georgia as far north as latitude 32° 45'. 

Hypotheses of the Formation of (^oral-Reefs. 

During the past few years elaborate reviews of theories of the forma- 
tion of coral reefs have been published by Prof. W. M. Davis, the 
larger of which are referred to in the footnote below.* These reviews 
are valuable in presenting most of the important coral-reef theories, 
as they are understood by a physiographer, who is convinced of the 
adequacy of the Darwinian hypothesis. Numbers of complex phe- 
nomena associated with coral reefs are not considered, and his pre- 
sentations are not in all respects satisfying. Prof. B. A. Daly has 
reviewed the literature on coral-reef theory, particularly from 
the standpoint of an adherent of the glacial-control hypothesis.' 
The literatiu-e on coral reefs is so enormous, that in the present paper 
consideration can be given only to certain papers that largely deal with 
coral-reef hypotheses or that contain information on areas herein dis- 
cussed. The limitations of space cause me to omit references to 
many papers of much merit. 

Three kinds of coral reefs are generally recognized, namely: (1) 
Fringing or shore reefs Which, as the name indicates, occur along the 
strand line; (2) barrier reefs which occur at variable distances 
off shore and have lagoons from 1 or 2 to as much as 30 of even 40 
fathoms in depth between them and the strand Une; (3) atolls, which 
are ring-like and inclose lagoons above whose surface no land-masses 
of importance protrude. 

As the relations of barrier reefs and atolls to the platforms above 
which they rise constitute in the opinion of geologists the essential 
part of the theory of che development of Recent reefs, the warfare 
of coral reef theory has been waged over the interpretation of these 
relations, which are the conditions of changing or changed position 
of the strand lino and the part played by reef-forming organisms 
as constructional agents. 

1. According to Darwin ^ and Dana,* corals first form a fringing 
reef off the sloping shore of a subsiding land area; the reef grows 

t Davis, W. 11, Dana's confirmation of Darwin's theory of ooral reefs, Amer. Joum. Sci., ser. 4, vol. 35, 
pp. 173-188, 1913; The home study of coral reefe, Amer. Oeogr. Soc. Bull., vol. 46, pp. 661-677, 641-664, 721-780, 
1914; A Shaler memorial study of ooral reefs, Amer. Joum. Sci., ser. 4, voL 40, pp. 2S3-271, 1015; Problems 
Associated with study of coral reefs, Sdentifio Mmithly, vol. 2, pp. 213-383, 479-501, 557-512, 1916. Also 
»Teral short articles in Nat. Acad. Sd. Proc., vols^ 1, 2, 1915-1917. 

s Daly, R. A., The glaciatooDtroI theory of ooral reefs, Amer. Acad. Arts and Set, vol. 51, pp. 157-251,1015. 

s Darwin, C. R., Structure and distribution of coral reefs, ed. 3, fig. 5, p. 134, fig. 6, p. 137, 1899. 

* Dana, J. D., Corals and coral reefs, ed. 3, pp. 263, 267, 1890. 



Digitized by VjOOQIC 



242 



BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



upward at such a rate that its top remains near the surface of the 
water and through retreat of the shore it is converted into a barrier. 
Continued subsidence, where the inclosed land area is an* inland, 
may result in the production of an atoU circumscribing a lagoon 
without any land mass projecting above the water level. But this 
is not all. The Darwinian hypothesis involves more than mere 




Fio. 4. COPT OF Darwin's fiqube illvstuatinq cokvebsion of a FRiNOiMa into a babrieb rbbf, 

ACCOBDINO to HIS HT70THBSIS. AA—OVtEVt EDGE OF TEE BEEF AT THE LEVEL OF THE SEA. BB— 
SHOiBSfl OF THE ISLAND. A'A'—OXTtEK EDOE OF THE BEEF, AFTER ITS UPWARD GROWTH DURINQ A 
PERIOD OF SUBSIDENCE. CC— THE LAOOON-CHANNEL BETWEEN THE REEF AW) THE SHORES OF THE 
NOW RNOBCLED LAND. B'B'—TWR SHOBBS OF THE ENCIRCLED ISLAND. N. B.— In THIS, AND THE 
TOLLOWINO CUT, THE SUBSIDENCE OF THE LAND COULD ONLY BE RBPRBSENTED BT AN APPARElTr RISE 
IN THE LEVEL OF THE SEA. 

subsidence and the conversion of a fringing into a barrier reef. It 
also attempts to account for extensive submarine platforms by 
assuming that they have been built upon sloping basements through 
agencies dependent on the presence of reefs. (See text-figs. 4, 5, 6.) 
Dana's interpretation " is e^entially that of Darwin. 




Fio. 5. Copt of Dabwin's hoube illustbatino convebsion of a babbibb beef into an atoll, ac. 
, ooRDiNO to bis htpothesis. A*A'—0\jteb xdqbs of the barribr-bbef at the level of the sea. 

The cocoa-nut trees represent coral-islbts formed on the beef. CO— The laooon-channel. 

B'£'— The shobes of the island, geneballt fobmed of low alluvial land and of coral detbi- 

TU8 FROM THE LAGOON CHANNEL. A"A"—TTtE OUTER EDGES OF THE REEF, NOW FQRMINO AN ATOLL. 

C— The lagoon of the newlt formed atoll. According to the scale the depth of the lagoon 

AND OF the lagoon CHANNEL IS EXAGGEBATED. 

That Darwin considered an alternative hypothesis is shown by the 
following quotation: . 

I may here observe that a bank either of rock or of hardened sediment, level with 
the surface of the sea and fringed with living coral, would be immediately converted 
into an atoll, without passing, as in the case of a reef fringing the shore of an island, 
through the intermediate form of a barrier reef. 



1 Corals and ooral islands, ed. 3, figs. pp. 263, 367, 1890. 



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GhBOLOOY AKD PALEONTOLOGY OF THE OAKAL ZONE. 248 

He adds, however, 

* * * but as we have seen, the larger groups of atolls in the Pacific and Indian 
Oceans have not been ionned on banks of this nature. ^ 




Fio. 6. RsFBODxrcnoN or J. B. Jukes' section across the great barrier reef of Austraua. a. 
Sea ouTsmE the barrier, generally unfathomable, b. The actual barrier, c. Clear chan- 
nel INSIDE the barrier, GENERALLY ABOUT 15 OB 20 FATHOMS DEEP. d. THE INNER REEF, f . SHOAL 
CHANNEL BETWEEN THE INNER REEF AND THE SHORE. F. THE GREAT BUTTRESS OF CALCAREOUS ROCK, 
FORMED OF CORAL AND THE DETRITUS OF CORALS AND SHELLS. G. THE MAINLAND, FORMED OF GRANITES 
AND OTHER SIMILAR ROCKS. 

2. The first important protest against the Darwinian explanation 
was by Carl Semper,' who, m 1863, after studies in the Pelew Islands, 
advanced the hypothesis that atolls could be formed in areas of 
elevation by the solution of the interior of preexistent limestone 
masses, and that solution, erosion by currents, and wave-cutting 
could develop platforms behind fringing reefs, thus transforming a 
fringing into a barrier reef. 

3. Murray* introduced the idea of banks being built upward 
by showers of the remains of pelagic organisms \mtil the bathymetric 
zone of reef-forming organisms is reached, and he called attention to 
the cutting of volcanic islands down to wave base. His theory has 
been briefly summarized by himself in the following words:* 

That when coral plantations build up from submarine banks they assume an atoil 
form, owing to the more abundant supply of food to the outer mar^, and the removal 
of dead coral rock from the interior portions by currents and by the action of the 
carbonic add gas dissolved in sea-water. 

That barrier reefis have been built out from the shore on a foundation of volcanic 
debris or on a talus of coral blocks, coral sediment, and pelagic shells, and the lagoon 
channel is formed in the same way as a lagoon. 

That it is not necessary to call in subsidence to explain any of the characteristic 
features of barrier reefs or atolls, and that all these features wo^ild exist alike in areas 
of slow elevation, of rest, or of slow subsidence. 

4. H. B. Guppy in 1890 published the following important opinion 
regarding the relations of barrier reefs to submarine plateaus or 
ledges:' 

I have now gone far enough to establish the probability, judging from the instance of 
the Australian Barrier-reef, that reefs of this class are in reality, and not in appearance, 

1 structure and distribution of ooral reefs, ed. 3, pp. 138, 139. 
< Semper, Carl, Reiaeberlcht, Zeitsch. fflr wiss. Zoologle, vol. 13, pp. 563-509, 1803. 
« Murray, John, On the structure and origin of coral reefs and islands, Roy. Soc. Edinburgh Proc., 
vol. 10, 1879-80, pp. 506-618, 1880. 
* Idem, p. 517. 
» Ouppy, H. B., The origin of coral reefs, Victoria Inst. Joum. Trans., vol. 23, pp. 51-61, 1800. 



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244 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

situated on the border of a submarine plateau or ledge. Such a position, according 
to the explanation of barrier-reefs, first advanced by LeConte, and supported by 
myself, presents the most favorable conditions for reef growth, the corals being limited 
on the outside by the depth, and on the inside by the sediment in the water. The 
influences of food-supply and currents act subsequently as auxiliary causes. 

What, then, is the explanation of the submarine ledge? The supposition that it is 
a continuation of the land slope is at once negatived by the fact that the slope of the 
land in the reef -encircled islands of the Pacific is usually 6 degrees or 7 degrees, some- 
times only 3 degrees or 4 degrees, but often as much as 10 degrees, or 12 degrees, whilst 
the submarine ledge, when stripped of reefs and defined by the 100-fathom line, would 
possess, a scarcely recognizable inclination, represented by a fraction of a degree. 
It will be found, however, when we examine the contour of such an island as Vanikoro, 
that the distance of the barrier-reef from the coaat may vary according to the slope of 
the land. Thus, on the west side of this island, the average angle of the land slope is 
6 degrees, and the distance of the barrier reef about 2^ miles. On the north side the 
inclination of the land is between 11 degrees and 12 degrees, and the barrier reef is 
rather over a mile distant. This is just what we should expect. The more gradual the 
land slope, the broader will be the submarine ledge, cut out in the course of ages by 
the action of the sea, and the more distant will be the barrier reef that has grown up 
along its margin. This I believe to be the true explanation of the position of barrier 
reefs. A submarine ledge is in the first place necessary; and, since the sediment and 
mud in the shallower waters on the ledge repress the growth of c(hu1s, reefs will 
naturally spring up toward the maigin of the ledge, where the water is clearer and 
where the depth is within that of the reef-coral zone.* 

5. Admiral Sir W. J. L. Whajton ' explained the uniform depth of 
atoll lagoons, whose edges are in various degress encircled by growing 
coral, by considering that the corals grow upon foundations that are 
the bases of volcanic islands that have been reduced by wave action 
to wave base. 

6. Alexander Agassiz ' foimd older limestone under the recent reefs 
in many areas investigated by him. He explained atolls by the solu- 
tion and erosion of the interior of preexisting limestone masses and 
ascribed the formation of the platforms of barrier reefs to marine 
erosion without change of sea level. 

7. Andrews ^ pointed out that the platform of the Great Barrier 
Reef of Australia has been submerged at a relatively recent date and 
that it continues southward beyond the reef, and he inferred that only 
a minor part of the platform is ''formed of coral growth." 

8. The opinions of Stanley Gardiner* are closely in accord with those 
of Semper, Murray, Wharton, and Agassiz. According to him sub- 
marine planation is effective to depths as great as 200 fathoms. 

1 Qappy, H. B., The origlii of ooral reefs, pp. SO, 61. 

* Wlu^toD, W. J. L., FoondfttiaDB of ooral atoUs, Nature, vol. 65, pp. 89(^-39S, 1807. 

* Agassis, Alexander, The Coral reefs of the Tropical Paoiflo, Mem. Mus. Comp. Zool., vol. 27, l toI. of 
text, 8 vols, of pis., lOOe. 

4 Andrews, E. C, Freliminary note on the geology of the Qoeenslaiid ooast with refereooes (to the 
geography of the Queensland and N. 8. Wales Plateau, Proc. Linn. 8oc. New South Wales, pt. 2, pp. 146- 
185, 1902. 

* Gardiner, J. Stanley, The Ibrmatioos of the Maldives, Qeographioal Journal, pp. 277-296» March, 1902; 
Fauna and geQgr^>hy of the Maldive and Laocadive Arohipelagoee, pp. 182, 183, 1901-3. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 245 

9. Hedley and Griffith Taylor * accepted Andrews's interpretations 
and clearly showed that coral reefs of either atoU or linear form that 
rise above shallow platforms owe their shapes to prevailing winds and 
currents. They say: 

This explanation differs from that of Sir J. Murray, who considers the atoll form to 
be aasnmed by abundant growth of well-fed corals on the maigin and the solution of 
dead coral rock in the interior. But if solution be so destructive, how can a reef form 
at all?' 

10. According to Daly' the depths in the drowned valleys within 
barrier reefs, in barrier-reef lagoons, and in atoll lagoons in the Pacific, 
are closely accordant and he attributes this accordance to Recent rise 
of sea level subsequent to deglaciation, whereby the depth of water 
in the Tropics was increased some 33 to 38 fathoms, thus submerging 
antecedent platforms of marine planation. That glaciation and 
deglaciation effect the development of living reefs did not originate 
with Daly, but it is principally he who has elaborated the hypothesis. 
He gives in his papers an accoimt of the earlier suggestions. 

11. Wood Jones* considered sedimentation the critical factor in 
coral-reef theory, as corals grow only where there is comparatively 
little deposition of sediment. He accepts the conclusions of Hedley 
and Griffith Taylor on the importance of winds and currents in shap- 
ing atolls, and especially attacks the hypothesis of ''a deepening or 
widening of the lagoon by a process of 'solution'." 

Although the results of my own investigations will be elaborated 
on subsequent pages, the following summary statement may here be 
made: I have greatly multiplied the evidence in favor of Recent 
submergence in the coral-reef areas in the western Atlantic, the Gulf of 
Mexico, and the Caribbean Sea, and have shown that the living off- 
shore reefs in those areas formed either during or after submergence 
and are growing on submerged basement platforms where conditions 
are favorable for the life of reef-forming corals. The platforms are 
continuous beyond the limits of the reefs and their existence is in no 
wise dependent upon the presence of reefs. 

I have also shown that the great Florida Plateau has existed as a 
plateau since at least late Eocene time; and that some of the West 
Indian platforms are about as old. As these plateaus existed previous 
to Pleistocene time they could not have been formed by marine plana- 
tion during Pleistocene glaciation. Whatever be the cause of shift 
in position of strand line, off-shore reefs form on shallow submarine 
flats during or after rise in sea level, provided the rate of movement 
be not too rapid. This explanation applies to the fossil reefs of 

I Hedtoj, C, and Taylor, T. Griffith, Coral reete of the Great Barrier, QneeoBlaDd, Atvtcalaeian A»oc. 
AdT. Sd., Adelaide Meeting;, pp. 897-418, 1907. 

t Idem., p. 407. 

* Daly, R. A., PMBtoewio glafllatfm and the ooral-reef proUem, Amar. Jouni. 8d., ter. 4, vol. 30, pp. 
297-306^ mO; The Qladal-ooDtrol theory of ooral reets, Amer. Acad. Arts and Sd., vol. 61, pp. 167-248, 1916. 

« Jooee, F. Wood, Corals and Atolls, London, 1910. 



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246 BULLBTIK 108, X7NITBD 8TATBB KAIIONAL MUfiBTTM. 

Florida and the West Indies as well as to the reefs living to-day. 
I have pointed out that there are in the Virgin and northern Leeward 
Islands and off the shores of Central America certain submarine 
terrace flats, one at a depth of about 17 to 20 fathoms, another at a 
depth of about 26 to 30 fathoms, the deeper flat being separated from 
the shallower by an escarpment. These relations accord with the 
demands of the Glacial-control theory as expounded by Daly. 

Tests of Coral-reef Htpotreses. 

The tests of the theories comprise ascertaining the answers to the 
following questions: 

1. Were the important coral reefs of the world formed during or 
after the submergence of their basements, either by a sinking of the 
land or by a rise of ocean level due to some world-wide cause ? 

2. What is the r61e of corals as constructional geologic agents i 
What percentage of the sediments around coral reefs is composed of 
corals, and is the flat area between a barrier reef and the shore due 
to infilling behind the reef or was there a shallow marginal flat before 
the reef formed t 

3. Can a lagoon channel behind a barrier reef or the lagoon within 
an atoll rim be formed by submarine solution by sea water or by 
submarine scour! 

4. Wbat and how much effect have wind-induced and other cur- 
rents in shaping coral reefs t 

5* What effect have glaciation and d^laciation had on the 
development of living coral reefs t 

Before considering the fossil and living coral reefs of the West 
Indies in their bearing on the answers to these questions, some of 
the more important criteria to be used in answering the questions 
will be briefly outlined. 

CRITERIA FOR RECOGNIZING SmTT IN THE POSITION OF STRAND LINE. 

The criteria for recognizing elevation of a former strand line 
comprise: (a) Coastal terraces bordered inland by escarpments or 
cliffs that may be inferred to owe their origin to wave cutting; 
(J) wave-cut grooves in cliffs and sea caves that stand too high to 
have been formed at present sea level; (c) elevated beaches or bars, 
which under proper conditions form on shallow marine terraces and 
at the mouths of embayments; (d) the presence above sea level of 
organisms that must have lived in the ocean. 

The criteria for recognizing submergence of former strand lines 
comprise: (a) Indwitation of the coast line caused by the sea invad- 
ing the lower parts of subaeriaUy eroded valleys, the channels of 
which in many instances are preserved below sea level across and 
beyond the existing strand line ; (b) the presence below sea level of 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 247 

submarine flats separated by relatively steep slopes or escarpments, that 
are due either to marginal wave cutting by the sea or are due to the 
formation of a subaqueous profile above a previous profile;^ (c) the 
presence, especially in limestones, below sea level, of solution wells, 
pits, and caverns that inferentiaUy were formed subaerially by the 
solvent action of fresh water; (d) the presence inland of free openings 
that connect with the sea, showing that there are underground 
channels by which ground water formerly flowed to the sea; (e) the 
presence of submerged peat bogs or swamp deposits composed of 
plants that grow only at or above sea level; (f) the presence below 
sea level of indurated limestone, the induration of which is due to 
solution of some of the original material and subsequent redeposi- 
tion f (g) erosion unconformities at the bases of marine formations, 
showing that there was subaerial erosion of the basement previous 
to the submei^ence during which the formation was deposited or 
accumulated in the sea. 

The foregoing statements might be elaborated, but to do so seems 
imnecessary. The criteria enumerated are those I have actually 
used in my own work. 

Besides ascertaining the proper succession of changes in the posi- 
tion of strand line, it is essential that the amount of the oscillations 
be measured, that differential crustal movements be noted and dated, 
and that an estimate be made of the endurance of the strand line in 
its relation to present sea level. 

CBITERIA FOR MEASURING THE AMOUNT OF VERTICAL SHIFT IN STRAND LINE, 
AND FOR DETERMININO THE RELATIVE AOES OF TERRACES AND THE PHT8I0- 
ORAPmC STAGE ATTAINED BT A SHORE LINE. 

The criteria for estimating the exact amount of rise or fall of sealevel 
are not yet definite, because adequate study has not been made of 
the factors that determine effective wave base and of the depth to 
which effective wave cutting extends. Notwithstanding this inade- 
quacy of precise information, an approximation of the amoimt of 
change may be made. In the case of elevation, the base of a wave- 
cut escarpment or cliff, the flats of marine terraces, and wave-cut 
grooves on sea cliffs, may be assumed to represent approximately 
former sea level. Approximate measures of the amount of sub- 
sidence may be based upon the depth of drowned valleys, the depth 
below sea level of the bottoms of submerged solution wells and 

> For dlseoaaiaiis of this subject see as Ibllows: 

Barrell, Joseph, Factors in movement of the strand line, Wa8hingtonAcad.Sd.Jocini.,vol.l3,pp.41S- 
420, 1915; Factors in movements of the strand line and their results in the Pleistocene and Post-Pleisto- 
cene, Amer. Joom. Sd., ser. 4, vol. 40, pp. 1-22, 1015. 

Vanghan, T. W., Some littoral and subttttoral physiographic features of the "^rgin and northern Lee- 
ward Islands and theirbearing on the coral reef problem, Washington Acad. Sd. Joum., vol. 0, pp. 
59-^lOie. 

> The fundamental principle of this criterion is discussed on p. 250, under the caption "SoHibility of cal- 
cium carbonate in sea water." 

37149--19--Bull. 103 5 



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248 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

cavemsy and the depth to which peat or swamp deposits that were 
formed at or above sea level are submerged. Where there are recog- 
nizable submerged wave-cut scarps, the depth of the base of the 
scarp below sea level is nearly a measure of the amount of sub- 
mergence; the depth in the West Indies in some instances probably 
exceeds the amoimt of submergence by about 6 fathoms. In the 
case of islands that rise from a common platform and which biologic 
and other data show were once parts of one land mass, the depth of 
water on the common platform may be assumed to be an approxi- 
mate measure of the amoimt of the rise of sea level with reference to 
those islands. 

The criteria for determining the relative ages of elevated terraces 
with reference to each other and for determining the amount of 
deformation to which they have been subjected are as follows: (a) 
Relative height; (h) relative amoimt of dissection; (c) relative degree 
of inclination and direction of the slope of the terrace flats; (d) pres- 
ence or absence of a succession of higher and lower terrace flats on 
promontory tips and in places protected from vigorous marine cut- 
ting; (e) stratigraphic relations of terrace deposits. 

Estimates of the endurance of the present relation of sea level 
to strand line are based upon recognizing the stage of |>hysiographic 
development of the shore line. Among the important features to 
be observed are the presence or absence and the character of sea 
cliffs bordering the shore; the amoimt of delta and alluvial plain 
building at the mouths of stream ways; the character of beaches, bars, 
and spits; the nature and extent of the alluvial deposits back from 
the shore; the profiles of valley sides; and the axial profiles of the 
streams. 

CRITERIA FOR ASCERTAINING THE r6lE OF CORALS AS CONSTRUCTIONAL 

AGENTS. 

The failure correctly to evaluate corals as geologic agents has been 
a defect of nearly all investigations of the so-called coral-reef prob- 
lem; in fact, usually no attempt has been made to make such an 
evaluation. This evaluation may be made in several ways, which 
are as follows: (a) In studying fossil reefs exposed to view, the 
relative proportion of coral to other constituents of the rock should 
be estimated; (h) in studying marine bottom samples, percentage 
estimates of the proportion of the different ingredients should be 
made; (c) for submerged platforms on which reefs grow, the area 
of the reefs should be compared with the total area of the platform, 
an effort should be made to ascertain the nature of the rock under- 
lying the sea floor between the reef and the shore, and the continuity 
in outline of the platform should be compared with the extent and 
position of the reefs; (d) knowledge of the growth rate of corals, 



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GEOLOGY AND PALEONTOLOGY^ OP THE CANAL ZONE. 249 

when the relative frequency of specimens is known, permits an 
estimate of the rate of their constructional work. 

This subject as a part of the problem of the formation of coral reefs 
possesses an importance that can scarcely be overestimated, for it 
comprises critical tests of both the Darwinian and the glacial-control 
hypotheses. The topics in the foregoing list will be discussed seridtim, 
(a) Estimate of the relative quantity of coral to other constituents 
in emerged formations containing reefs, if they have not been exten- 
sively recrystallized, is relatively simple, although great precision in 
quantitative expression is not to be expected. This topic will be 
further considered in discussing the Caribbean, floridian, and Baha- 
man fossil reefs. 

(6) Percentile estimates of material according to source are 
difficult, but the results are of great value. The technique of making 
such estimates is described in a memoir recently pubUshed by the 
Cam^e Institution of Washington.^ 

(c) Here it should be emphasized that one of the postulates of the 
Darwinian hypothesis is that the prism of material included between 
three surfaces, namely, (1) the sea-bottom landward of the barrier, 
(2) a surface assumed as an extension of the land slope under sea imtil 
it intersects (3) a surface projected downward from the landward 
face of the reef, is due to the presence of the reef (see figure 4, page 
242). Proof that a barrier has formed during or after submergence 
does not carry with it proof that the prism of material above indi- 
cated is due to the presence of the reef. 

There are at least three criteria that can be applied in deciding 
whether or no the flat between the reef and the shore exists inde- 
pendently of the reef. They are as follows: (1) If the flat is de- 
poident on the presence of the reef, where there are breaks in the 
barrier tongues of deep water should extend landward across the 
shaBow bottom of the flat behind the reef; and where there is no reef 
there should be either a normal profile of equilibrimn or an approach 
to such a profile, showing a deeper flat than that behind the reef, be- 
cause of the absence of an off-shore wall behind which sediment 
could accumulate; but if the flat is independent of the reef, in general 
it should be continuous irrespective of the presence of the reef and 
should in places extend beyond the reef Umits. (2) If the formation 
of the flat is dependent on the presence of the reef, the reef should 
stand on the seaward edge of the flat, that is, the flat should not project 
seaward beyond the reef. (3) It is often possible to discover the 
nature of the rock forming the sea floor between a barrier and the 

1 vaughao, T. W., in ooUaborstlon with Cuahman, J. A., Ooldman, M. I., Howe, 11 A., and others. 
Some shoal-water bottom samples from Murray Island, Australia, and comparisons of them with samples 
from Florida and the Bahamas, Carnegie Inst. Washington Pub. 213, pp. 235-2»7, pis. 94-98, 1918. See 
Qspecbaiy the article by M. I. Goldman, Composition of two Murray Island samples according to source of 
material' pp. 249-262. 



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250 BULLETIN 103, UNITEl/ STATES NATIONAL MUSEUM. 

shore. Such a floor if formed by agencies associated with the pres- 
ence of the reef will not be composed of rock demonstrably older 
than the reef, and will not exhibit geologic phenomena that in age 
clearly antedate the reef; but if it can be shown that the rock of the 
floor is older than the reef and that the floor has had a geologic his- 
tory antecedent to the formation of the reef, it is demonstrated that 
the reef is merely growing on the surface of a flat whose formation 
is entirely independent of the reef development. 

(d) The growth rate of corals, which furnishes one of the checks 
to be appUed to the Glacial-control hypothesb of the formation of 
living reefs, is further considered on pages 253, 254. 

SOLUBILITY OF CALCIUM CARBONATE IN SEA WATER. 

As the formation of lagoon channels behind barrier reefs and of 
atoll lagoons by the solvent action of carbon dioxide (COj) dissolved 
in sea water is a part of the coral-reef hypotheses of Semper, Murray, 
A. Aggassiz, and Gardiner, if lagoons and lagoon channels have hem 
formed in the way indicated, in the Tropics the surface waters of 
the ocean should contain an excess of carbon dioxide (OQi) and 
should exercise a demonstrable solvent effect on calcium carbonate 
(CaCOs). If it should be foimd that there is no excess of carbon 
dioxide (CO,) in such water and that the water is saturated with 
reference to calcium carbonate (CaCOs), the hypothesis of the forma- 
tion of lagoons and lagoon channels in the manner postulated by 
Murray and 'others must be definitely abandoned. 

In 1913, Mr. K,. B. Dole undertook at Tortugas, Florida, certain 
examinations that were intended to solve this problem, if possible. 
In 1914 I summarized in the following words the results I had ob- 
tained from a study of the bottom samples along the Florida reef 
tract, those of Drew on dentrifying bacteria, and those of Dole on 
the chemistry of the waters.^ 

There are two rival hypotheses for the formation of atolls: One of these attributes 
them to the submarine solution of the interior of a mass of limestone, the other ac- 
coujits ior them by constructional agencies. In order thorougjily to test the solution 
hypothesis the results of four lines of investigation were brought to bear upon it, 
and all are accordant. (1) All the bays, sounds, and lagoons within the Florida reef 
and key region are filling with sediment; (2) Drew's investigations of dentrifying 
bacteria show that chemical precipitation of calcium carbonate is taking place in 
the lagoons; (3) the chemical examination by R. B. Dole of samples of sea water 
flowing into and out of Tortugas lagoon, collected twice daily for a lunar period, show 
that although both carbonate and bicarbonate radicles are in solution uncombined 
carbon dioxide is not present, and that the water possesses no capacity for further 
solution of calcium carbonate by virtue of its content of free carbon dioxide; (4) the 
determinations by Dole of the salinity of the water within the T<»rtuga8 lagoon and 
at the southern end of Biscayne Bay show a higher concentration than that in the 
open sea water on the outside, indicating that tidal inflow and outflow are not suffi- 
cient completely to mix the water in the lagoons with the water of the surrounding 

1 Wash. Acad. 8d. Joam., yoL 4, pp. 27-38^ Jan. 19, 1914. 



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GEOLOGY AND PALEONTOLOGY OP THE OANAL ZOKE. 251 

sea and that concentratioQ by evapcmttion is taking place. As the results of these 
lines of inquiry are so positive, the formation of lagoons by submarine solution may 
be definitely eliminated from consideration. 

Since the publication of this statement other investigators have 
made important contributions to this subject, noteworthy among 
whom are John Johnston, H. E. Merwin, and E. D. Williamson, of 
the Geophysical Laboratory of the Carnegie Institution of Wash- 
ington, and Roger C. Wells, of the United States Geological Survey. 
Wells says: * 

In other words, sea water [from the Florida reef] appears to contain so much car- 
bonate that in contact with the atmosphere at 1*^ C. it neither has nor acquires an 
appreciable solvent action on caldte. 

As I have considered the subject in detail in my paper on the 
Murray Island bottom-samples' and in a paper on "Chemical and 
oi^anic deposits of the sea" * I will merely say that sea water in 
shoal-water areas within the Tropics can not dissolve calcium car- 
bonate, and that lagoon channels and atoll lagoons are not formed 
by solution, but are flattish areas more or less completely inclosed by 
built-up walls. 

As lagoons are areas of sedimentation and not of removal of 
material, their formation by submarine scour may also be discarded. 

EFFECTS OF WIND-INDUCED AND OTHER CURRENTS IN SHAPING CORAL REEFS. 

This is an old topic; in fact, considerable bibliographic work 
would be needed to ascertain the names of all the investigators who 
have contributed to it and who deserve mention. That Darwin at 
least had an adumbration of the importance of these agents is indi- 
cated by his statement regarding Keeling atoll: ^ 

That they [the waves] beat against it in the same peculiar manner in which the sweU 
from windward now obliquely curls round the margin of the reef, was evident from the 
conglomerate having been worn in to a point projecting from the beach in a similarly 
oWque manner. 

Among recent investigators Hedley and Griffith Taylor, as noted 
on page 245, Wood Jones,*^ and I, in a number of my papers, two 
of which are cited below,' have devoted attention to this subject. 
During the field season of 1914 I had numbers of Ekman meter 
current-measurements made around Tortugas and at other places 
along the Florida reef tract. The measurements to a certain degree 

> Wells, R. C, The solubility of caldte in sea water in coDtact with the atmosphere, and its variation 
with temperature, Carnegie Inst. Washington Pab. 213, pp. 316-318, 1018. 
a Carnegie Inst. Washington Pub. 213, pp. 205-288, 1917. 

• QeoL Soc. Amer. BulL, vol. 28, pp. 933-944, 1918. 

• Stru c tur e and distribution of coral reefs, ed. 3, p. 22, 1889. 

• Coral and atolls, pp. 253-261, 1910. 

• The buUdtng of the Marquesas and Tortugas atolls and asketchof the geologic history of the Florida reef 
traet, Carnegie Inst. Washington Pub. 182, pp. 56-67, 1914; Sketch of geologic history of the Florida coral - 
reef tiact and comparisons with other coral-reef areas, Washbigton Acad. Sci. Joum., vol. 4, pp. 26-34, 
1914. 



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252 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

give qualitatively the relations of currents to land forms, and com- 
pletely confirm the more qualitative generalizations of Hedley and 
Griffith Taylor, which in brief are the axis of elongation of linear reefs 
is parallel to the direction of the dominant current while the bow of 
a crescentic reef is directed toward the direction whence the dominant 
current comes. These relations of reef form to current direction are 
most striking where the reefs rise above comparatively shallow plat- 
forms, as along the Great Barrier reef of Australia and along the 
Florida Keys. In atolls that more or less encircle the flat tope of 
submarine peaks, although currents are imdeniably important in 
shaping sections of the reefs, they are not of so great importance as 
reefs that rise above shallow, long, wide platforms. 

CRITERIA FOR DETERMINING THE EFFECT OF GLACIATION AND DEGLACIATION 
ON THE DEVELOPMENT OF LIVING REEFS. 

Daly's elaborate paper on the Glacial-control theory of coral reefs 
has been cited on page 245. If the Glacial-control theory is true the 
following conditions should prevail: (a) There should be evidence of 
geologically Recent submergence of most of the shore-lines of the 
earth; (h) the average amount of submergence should be equal to the 
amount of lowering of the ocean-level during Pleistocene glaciation; 

(c) the position of the strand line during Pleistocene glaciation should 
be indicated by scarps separating flats, and the amount of sub- 
mergence indicated by their present position below sea level should 
agree. with the amount of raising ocean level due to deglaciation; 

(d) rate of growth corals should be such that since the disappearance^ 
of the continental ice sheets coral reefs could grow to a thickness 
equal to the amount sea level was raised as a result of the deglaciation; 

(e) living barrier coral reefs and atoll reefs should be superposed on 
antecedent basement flats or platforms. It should here be stated 
that the fact that there has been local differential crustal movements 
does not at all invaUdate the importance of the Glacial-control theory 
in its application to the explanation of the modem coral-reef develop- 
ment. 

Of the criteria stated in the foregoing list only the amount of 
vertical change in the position of sea level because of glaciation and 
deglaciation, the length of time since the disappearance of the great 
continental glaciers, and the rate of growth of corals need discussion 
at this place. After their consideration some attention will be given 
to other criteria of less determined value. 

AMOUNT OP VEBTICAL DISPLACEMENT Or STBAND LINE BY QLAOATION AND DBOLAOATION. 

It is entirely obvious that the withdrawal of water from the ocean 
to form the Pleistocene continental glaciers would lower sea level, 
and that the return of the waters so locked up to the ocean upon the 
melting of the continental glaciers would raise sea level back to where 



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GEOLOGY AND PALEONTOLOGY OP THE OAFAL ZONE. 253 

it stood previous to the formation of the continental glaciers, unless 
crustal changes in the earth counterbalanced the effects of such with- 
drawal and return of oceanic water. Reference here will be made 
to only the two latest computations. 

W. J. Humphreys, as part of a symposium before the Geological 
Society of Washington, on March 24, 1915, said:^ 

The fact that the average thickness of the ice cap during the last glaciation can be 
only roughly estimated renders any calculation of its effect on ocean level corre- 
spondingly doubtful. It does not seem probable, however, that they should have 
averaged much if any thicker than the present caps of Greenland and of Antarctica, 
which a number of good observers have estimated to be about 1,000 meters. Taking 
this value and assuming the d^ladated area to be equal to one-fifteenth the area of 
the ocean, or, roughly, twice the glaciated area of North America, we estimate the 
change in sea level to have been about 67 meters. As already stated, this is only 
an estimated change, but perhajM it is a conservative estimate. 

Daly in his paper on the Glacial-control theory of coral reefs sum- 
marizes his discussion in the following words:* 

Combining results, it is seen that, at the time of maximum glaciation, the tropical 
seas probably had an average level which was 60 to 70 meters (33 to 38 fathoms) lower 
than at the present time. 

The estimates of Humphreys and Daly are essentially the same. 

As maximum glaciation was probably not of long duration the 
greatest effect of submarine terracing would be expected in some- 
what shallower depths, probably between 20 and 30 fathoms. 

BATE or OBOWTH OF COBALS AND LBNGTH OF POOT-OLACIAL TIME. 

Recently I have published two summaries of the results of my 
experiments and observations on the growth rate of Floridian and 
Bahamian corals, and compared my results with those obtained by 
investigators in the Pacific' The following statements are taken 
from the second of the papers referred to in the footnote: 

As has been stated, the primary object of this investigation was to get an approxi- 
mate measure of the rate at which corals might build reefe. In order to make this 
estimate the true reef corals must be considered separately from those which live in 
other habitats. The reef species jHtr excellence in the Recent and Pleistocene reefe of 
Florida and the West Indies is Orbicdla annularis; after it in importance are Maean- 
dra 9trigo$a, M, Idbyrinthtformis, and Siderastrea siderea. Other corals, the most impor- 
tant of which is Pontes astreoides, with Agarida and Favia fragum of secondary 
importance, occur in the areas intermediate between the prominent heads. In some 
areas Acropora palmata ia the dominant species. The massive heads fonn the strong 
framework of the reef, with infilling by other corals and other oiganisms. Therefore 
the upward growth rate of Orbkella anmUaris on the reef is critical. * * * 

1 Htimphre>s, W. J., Changes of sea lev^l due to changes of ocean vohime, Washington Acad. Bd. 
Jonm., vol. 6, pp. 446-446, June 19, 1916. 

< Amer. Acad. Arts and ScL Proo., vol. 61, p. 174. 

« Vaughan, T. W., Geologic significance of the growth-rate of the Floridian and Bahaman shoal-water 
oorab, Washington Acad. Sd. Joum., vol. 5, pp. 501-600, 1915; Growth rate of the Flondian and Bahaman 
shoal-water corals, m On Recent Madreporaria of Florida, the Bahamas, and the West Indies, etc., 
Carnegie Inst. Washington Yearbook No. 14, pp. 221-231, 1916. 



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254 BULLETIN 103, UNITED STATES NATIONAL MUSEX7M. 

Uaiiig these figures [in the paper referred to] as the basis of a further computation, 
a reef by the continuous upward growth of c<»rals [Orbicella annularis] might attain 
at a rate of 6 mm. a year a thickness of 25 fathoms«150 feet in 7,620 yean; and at a 
rate of 7 mm. a year it might attain the same thickness in 6,531 years. 

Should the growth rate of Acropora pcUmata be taken as a measure, the time to 
accumulate such a thickness would be considerably less. This species formB spread- 
ing, palmate fronds, rising from stout bases. As age advances the fronds thicken and 
can withstand the pounding of surf and breakers. The average upward growth is 
between 25 and 40 mm. per year, but as the interspaces between the fronds are con- 
siderable in volume, comparisons with Orbicella anntUaris must be based upon relative 
increases in weight for a known period. * * * 

These two estimates [as shown in the paper dted] give a measure of the limits d 
reef formation under continuously favorable conditions for upward growth. Such 
corals as Orbicella annulariM might form a reef 150 feet thick in between 6,500 yeaiB 
and 7,600 years; while such xx>rals as Acropora palmata might form a similar thickness 
in 1,800 yean. 

«»»♦♦»♦ 

The data available for the Pacific corals are not so abundant as those for the Atlantic, 
nor have the records, with few exceptions, the same degree of precisbn. However, 
they are siifiicient for some general comparisons. The general growth rate of branching 
corals is nearly the same for both regions; but the growth of the massive forms in the 
Pacific appean to be appreciably more rapid than that of similar forms in the Atlan- 
tic. Therefore it seems probable that in the coral reef regions of the Pacific and Indian 
oceans a reef 150 feet thick may form under favorable conditions in less than 6,000 
yean. According to Gardiner such a reef mi^t form in 1,000 yean. 

As the disappearance of the last continental ice sheets is estimated to have been 
between 10,000 yean ago in Scandinavia and Alaska and 40,000 yean ago at Niagara, 
the data presented show that there has been ample time tat the developm^it of any 
known living reef since d^ladation. 

imoT Of LowsBoro or uaboxe tbmpkbatube on bekt oobais Dumoro ouLCunoir. 

Daly in his paper on the Glacial-control theory devotes much atten- 
tion to the probable extinction of reef corak over large areas and 
their restriction to only the hotter parts of the ocean during glacia- 
tion.^ Daly's discussion of this subject is interesting and suggestive, 
but not really convincing. It is one on which far more research is 
needed. I rather hope that the data I have recently presented in 
my paper on the temperature of the Florida coral-reef tract* will aid 
in furnishing a basis for such a computation. That there was a 
lowering of the vitahty of corals over large areas marginal to tropics 
can scarcely be doubted, but that reef corab thrived throughout 
Pleistocene time appears more than merely probable. 

In this connection this following list of corals from the elevated reefs 
of Barbados is pertinent. Professor Jukes-Browne sent the collection 
to me after Prof. J.W. Gregory had published his paper on the Bar- 
badian elevated-reef corals,' making the statement that great care 
had been taken in determining the height above sea level at which 

1 Amer. A.cad. Arts and Sd. Proo., vol. 51, pp. 16&-171. 

• Carnegie Inst. Washington Pab. 213, pp. 819-^39, 1918. 

* Gregory, J. W., Contributions to the paleontology and physical geography oC the West Indies, Oeol. 
Soc London Joum., vol. U, pp. 255-310, pL 11, 1305. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 266 

each lot was obtained. The collection is now the property of the 
United States National Museum. 

Corals from the elevated reefs of Barbados submitted by Prof A. J. JvJces- 

Browne, 

Elevation 1,043 feet. Horse Hill, St. Joseph. 

OrbiceiBa annularis (EUis and Solander). 
Elevation 845 feet. Cutting side of road, Parris Hill, St. Joseph. 

OrbiceUa annularis (EUia and Solander). 
Elevation 747 feet. Cutting side of road, Market Hill, St. George. 

OrbiceUa anrndaris (Ellis and Solander). 
Elevation 720 feet. Russia Gully, St. Thomas. 

OrbiceUa annularis (Ellis and Solander). 

Ma^andra hhyrintMforrnis (Linnaeus). 
Elevation 707 feet. Haynesfield, St. John. 

Stephanocoenia intersepta (Esper) 

OrbiceUa annularis (Ellis and Solander). 

lianidna gyrosa (EUis and Solander). 
Elevation 700 feet. St. Johns Church, St. John. 

Maeandra strigosa (Dana). 
Elevation 480 feet. Locust Hall, St. (jeorge. 

Stephanocoenia intersepta (Esper). 

OrbiceUa annularis (EUis and Solander). 
cavernosa (Linnaeus). 

Siderastrea siderea (Ellis and Solander). 
Elevation 362 feet. Ridge, Christ Church. 

Siderastrea siderea (EUis and Solander). 
Elevation 360 feet. SmaU Ridge, Christ Church. 

OrbiceUa annularis (Ellis and Solander). 
Elevation 300 feet. Skeens HiU, near Lower Greys, Christ Church. 

OrbiceUa anmilaris (EUlis and Solander). 

Siderastrea siderea (Ellis and Solander). 
Elevation 300 feet. Dayrells Ej31, St. Michael. 

Manidna gyrosa (EUis and Solander). 
Elevation 180 feet. Codrington Quarry, St*. Michael. 

OrbiceUa annularis (Elllis and Solander). 

Manidna gyrosa (Ellis and Solander). 
Elevation 160 feet. Cutting side of road, Charles Rose guUy, St. 
George. 

Maeandra Tahyrinthiformis (Linnaeus). 
Elevation 100 feet. Chelston Quarry, St. Michael. 

Meandrina maeandrites (Linnaeus). 

Manidna gyrosa (EUis and Solander). 

Siderastrea siderea (EUis and Solander). 

Acropora muricata (Linnaeus). • 



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256 BLLLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Elevation 80 feet. Prospect, St. James. 
SWphanocoenia intersepta (Esper). 
OrhiceUa annularis (Ellis and Solander). 
Maeandra IdbynnlMfarmis (Linnaeus). 
Acropora muricata (Linnaeus) s. s. (as pebbles). 
Elevation 70 feet. Grazettes, St. Michael. 
StepJianocoenia intersepta (Esper). 
OrhiceUa annularis (Ellis and Solander). 
Maeandra hhyrinthiformis (Linnaeus). 
Siderastrea siderea (Ellis and Solander). 
Elevation 40 feet. Sandy Lane^ St. James. 
OrhiceUa annularis (Ellis and Solander). 
Maeandra lahyrinthiformis (Linnaeus). 
Elevation 40 feet. Colleton, St. Lucy Parish. 

Maeandra strigasa (Dana). 
Elevation 20 feet. Black Rock. 

Acropora muricata (Linnaeus) s. s. 
Just how much of Pleistocene time is represented by this collection 
I can not say, but it is certainly a considerable part of it. 

Mr. O. E. Meinzer, in the vicinity of Guantanamo Bay, Cuba, ob- 
tained living species of reef corals on Pleistocene terraces between 400 
and 500 feet, at 275 feet, 200 feet, 125 feet, and 50 feet above sea level. 
It is unfortunate that Daly should have attempted to account for 
the disappearance in the West Indies of so large a percentage of genera 
that now persist in the Indo-Pacific by appeal to the lowering of the 
temperature in the western Atlantic Ocean through Pleistocene glaci- 
ation. In a recently published paper ^ as well as the present one, I 
have shown that the genera had disappeared previous to Pliocene time. 
It is at present my opinion that not enough is known regarding the 
effect of lowering of marine temperature during glaciation to serve 
as a basis for very strong arguments for or against the validity of the 
Glacial-control hypothesis. 

YALLBT-IN-VALLBT ABRANOBMBNT Ain> CUTTEO SPURS. 

Professor Davis says in his Shaler Memorial study of coral reefs: 

Furthermore, if the embayments of a central iflland within a barrier reef reeult from 
the drowning of valleys that were eroded with respect to lowered sea level of a relatively 
short glacial period, then each valley must be entrenched in the floor of a pregladal 
valley; and above the head of each embayment resulting from the drowning of a 
new-cut valley, there should be a 'Walley-in-valley'' landscape, unless the pre- 
gladal valley was so young and narrow that its sides were undercut and destroyed by 
the deepening and widening of the glacial valley.^ 

I Vaaghan, T. W., Tbe reef-coral Haona of Carrixo Creek, Imperial County, California, and its signiflcaace, 
U. 8. Geol. Surv. Prof. Pap. W-T, p. 366, 1917. 
« Amer. Joura. Scl., ser. 4, vol. 35, p. 240, 1915 



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GEOLOGY AND PALEONTOIXKJY OF THE CANAL ZONE. 257 

The character of the entrenching within an estabUshed valley after 
lowering of sea level will depend upon the off shore slope of the sea 
bottom previous to the lowering of sea level. As any acceleration 
of headward erosion by a stream depends upon increase in steepness 
of the longitudinal profile of the stream bed, unless the gradient of 
the lower course of the stream is considerably increased there will 
be no visible valley-in-valley landscape after submei^ence following 
d^laciation. Subsequently I will show that in the West Indies there 
is abimdant evidence of another kind that during Pleistocene time 
sea level was lowered, and that at the close of Pleistocene time it was 
raised. Valley-in-valley arangement is a criterion of very doubtful 
value. 

Professor Davis also insists that if the Glacial-control hypothesis 
is correct, the spurs of islands within barrier reefs should be cliffed — 
the cliffs cut during Pleistocene glaciation. As promulgated in print 
by Professor Davis, I doubt the vaUdity of this criterion. Perhaps 
the following hypothetical explanation may apply in some instances: 

Around volcanic islands, the centers of which are far enough from 
the shore for the surface profile of the eject a to have assumed the 
theoretic catenary curve, marine planation may proceed without at 
first cutting pronounced cliffs. If the material on the higher slopes 
is not greatly consolidated, alluviation and surface creep may deliver 
detritus more rapidly than the sea can remove it by mai^inal cutting 
and by undertow and other transporting agents. The sea may thus 
be held back from the interiorly situated harder volcanic rocks and 
the development of well-marked sea cliffs may thereby be prevented 
while the sea bottom would be aggraded near shore and a submarine 
flat produced. Should sea level then fall so that the shore line would 
shift to the outer edge of the previously formed flat, erosional processes 
might obliterate the low scarp carved into unconsolidated coUuvial 
and alluvial material. Under such circumstances, should the sea- 
bottom gradient be less than that of the stream profiles^ the lowering 
of sea level would not lead to the development of valleys-within- 
valleys, and alluvial plains might be pushed forward beyond the ends 
of the interstream spurs. Should sea level rise back to its former 
stand reef corals might establish themselves on the submerged flat 
at any place where the proper ecologic conditions might be found 
and develop into a barrier reef, off a land area on which there would 
be no valley-in-valley arrangement of stream courses and along whose 
shores there would be no cliffed spurs. This is an hypothetical in- 
stance, but that it is possible is apparently shown by the island of 
St. Christopher, West Indies, where such an arrangement of central 
volcanic mountains and relatively flat areas underlain by volcanic 
ejecta and coUuvial and alluvial material intervene between them 



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258 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

and the shore. In other volcanic islands the sea may not be held 
back from the harder rocks and may cliff them. 

There are numbers of possibilities which deserve consideration, but 
the actual explanation of how present conditions were brought about 
is possible only through detailed field work in each area. 

Some other kinds of shore lines may be mentioned. It is well 
known that one of the important factors in determining the amount 
of cliffing and the character of the cliffing of some shores is geologic 
structure. In an uplifted island composed of bedded sediments 
which have been moderately tilted the highest cliffs will be on the 
up-dip side along the line of the strike; the cliffs will decrease in 
height from the up-dip exposure along the line of the dip, and on the 
side of the island where the rocks pass beneath sea level there may be 
almost no cliffs. These relations are well illustrated in AnguiUa and 
other islands in the West Indies. After such an earth block has 
been outlined there may be oscillation of strand line without further 
local crustal deformation. 

The island of St. Croix is interesting in this connection. Just 
south of its north shore, which is determined by a fault, are maturdy 
dissected mountains which attain an altitude of about 1,000 feet. 
Off the south foot of the highland is a sloping, slightly undulating 
plain, underlain by limestone, which extends to the south coast, 
(See pi. 70, fig. D). If this island were submerged 120 feet the lime- 
stone plain would form a submarine flat from one to about three 
sea-miles wide. Corals might grow on such a flat and form a barrier 
reef inside which there would be no strongly cliffed spurs along the 
shore, while the moimtains would be in a stage of mature dissection. 

Ambrican Tbbtiart and Pleistocene Reef Corals and Coral Rbefs. 

Most investigators of the genesis of coral reefs have considered 
only the modem; but the ancient, or fossil, reefs in many instances 
afford better opportunities than the living reefs to determine the 
geologic character of the basement on which the reefs have been 
built, the change in the relation between the reef basement and sea 
level, and the importance of corals as constructional agents. The 
southeastern United States and near-by West Indian Islands furnish 
numerous examples of both ancient and modem coral reefs, and 
these have been the subject of investigation for many years. The 
location of the Tertiary fossil reefs in the southeastern United States, 
their associated faimas, the inclosing sediments, including in most 
instances both the overlying and imderlying strata, the stratigraphic 
relations of the successive geologic formations, the geologic structure, 
and the geologic history, have been ascertained with a fair degree 
of accuracy. The coralUf erous beds range in age from the base of the 
Eocene to Recent, and the coral fauna of each geologic formation is 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 259 

known with approximate completeness. The total coral faunas have 
yielded some hundreds of species. 

EOOENB KEEP CORALS OF ST. BARTHOLOMEW. 

The corals obtained from the St. Bartholomew limestone are listed 
on page 194. Although there are many specimens and species of reef 
f acieSy they scarcely form a reef properly speaking. However, the 
stratigraphic relations are interesting. The best collecting groimd 
is on the northeast face of the northwestward projecting limb of the 
island, between Anse L6zard at the northwest and Jean Bay at the 
southeast. Anse £caille lies between the two bays mentioned. 
Qeve's ^ accoimt of the geologic succession is correct, perhaps with 
some modification of his dates of a part of the igneous rocks. The 
base of the section is composed of volcanic agglomerate, above 
which there is interbedded agglomerate or sandstone, conglomerate 
composed of volcanic material, and limestone, succeeded by mas- 
sive, hard, blue limestone. Most of the corals occur in the lower 
part of the sedimentary formation, in the limestone or in the 
softer, more rapidly weathering layers of calcareous sandstone, in 
which there is rehandled volcanic material. In conglomerate at 
the base of one exposure I observed boulders of volcanic material 
as much as 8 inches in diameter. Although, as Qeve stated, there is 
some interbedding of the limestone and agglomerate in the lower part 
of the sediments the upper formation rests imconformably on the 
lower. 

The gradation upward into piurer, more massive limestone has been 
mentioned. The presence in the higher limestone of a few corals of 
the same species as those in the lower beds and the abundance of 
calcareous algae in some places, indicate a shoal-water deposit; 
and, as the area of the deposit is relatively extensive, the evidence 
is in favor of its having been laid down on a submerged flat. 

The Jamaican Eocene corals are shoal-water forms but they are 
really not of reef facies. 

WEST INDUN MIDDLE OLIOOCENE REEFS. 
ANnauA. 

That the bedded volcanic tuffs imderlying most of the Central 
Plain of Antigua dip imder the Antigua formation toward the north- 
east is indicated by the general structure of the island, and is con- 
firmed by a well record, kindly furnished me by Dr. H. A. Tempany, 
government chemist of the Leeward Islands. The record mentioned 
is of a well bored on Fitches Creek, half a mile northeast of the south- 
west boundary of the limestone. Compact, noncalcareous rock 
was struck below the limestone. In the Central Plain patches of 

> Cleve, P. T., On the geology of the northeastern West India Islands, K. svenska Vet.-Akad. Handl. 
vol. 9, No. 12, pp. 24-37, 1872. 



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260 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

gravel and cobbles overlie the surface of the bedded tuffs at a number 
of places, two of which are Casada Gardens and Gunthorpe sugar- 
factory. At Morris Looby Hill, near the head of WiUoughby Bay, 
conglomerate immediately underlies the limestone; and the basal 
contact of the formation is also exposed on the north side of Wil- 
loughby Bay, where it is underlain by conglomerate, mostly composed 
of basic volcanic material. The main reef occurs within the Antigua 
formation at or near its base and is exposed along a southwest- 
northeast line from WiUoughby Bay to near Wetherell Point. The 
Antigua reef therefore grew upon a basement that had been sub- 
aerially eroded and was later depressed below sea level. The reef 
and the limestone of which the reef forms a part were formed during 
or after the submergence of their basement. Associated with the 
corals are many specimens of several species of Lepidocydina, which 
are oi^anisms characteristic of shallow, tropical water. The areal 
extent of these sediments, coupled with the fact that the deformation 
of the water-bedded tuffs that lie below the Antigua formation is not 
much greater than that of the Antigua formation, indicates that 
they were deposited on a submarine flat. In the northeastern part 
of the island both the tuffs and the limestone, according to J. W. 
Spencer, dip northeastward at a rate of 12® to 20®.* My own meas- 
urements show dips of about 20® toward the north or northeast for 
the volcanic tuffs and dips between 10® and 15® in amount, and rang- 
ing from N. 60® E. to N. 70® E. in direction, for the Antigua formation. 
The rocks are more disturbed in the Central Plain, where the dips 
of the volcanic tuffs were measured. Therefore, according to the 
available evidence the Antigua formation was a relatively extensive 
formation deposited in shoal water on a flattish floor. 

The main reef-coral bed is about 60 feet thick and is near the 
bottom of the formation. Above it corals are scarcer, but appear 
to be too sparingly distributed throughout a thickness of about 300 
feet of limestone above tlieir profuse development nearer the base, 
or the Antigua formation seems to have a total thickness of a little 
more than 350 feet. 

PORTO RICO. 

The middle Oligocene coral faima, as has been stated on page 204, 
occurs in the geologic formation to which Hill applied the name 
Pepino. This is a hard, calcareous marl, full of coral heads, with 
occasional indurated strata of white porous limestone. It is weD 
exposed north and northwest of Lares in the Pepino Hills, whence 
the name for the formation is derived and where the coflection of 
corals submitted to me by Mr. Hill was obtained.' This is the for- 

1 Spencer, J. W., On the geological and physical development of Antigua, Oeol. Soc. London Quart. 
Journ., vol. 57, pp. 494, 496, 1901. 

s Hill, R. T., Notes on the forest conditions of Porto Rico, U S. Department of Agriculture, Div. of 
Forestry Bull. No. 25, pp. 14, 15, 1899. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 261 

mation to which C. P. Berkey later applied the name Arecibo forma- 
tion.^ I have no field acquaintance with the formation but from 
some notes on it made by Oeve, before it had been named , from the 
descriptions of Hill and Berkey, and from some of the corals coUected 
by members of the New York Academy Porto Rican investigations, 
I am inclined to the opinion, that there is not a ''formation" but a 
group of formations of similar lithology, for the ''formation" contains 
both middle Oligocene (Antiguan) and upper Oligocene (Anguillan) 
fossils, and probably also some Miocene species. Ultimately the 
"formation," as Berkey also has suggested, will probably be spht 
into several formations; it seems to me that there will be at least 
three and perhaps more. Only the stratigraphic relations of the base 
of the formation particularly need consideration here. These rela- 
tions are those of unconformity according to Berkey, who says:' 

Above it [the Arecibo fonnation] in all cases lie the recent alluvial deposits and tho 
San Juan formation, and below it lie the older and more complicated igneous and 
sedimentary rocks. The break between these two represents the chief unconformity 
in the whole geological column. 

An excellent illustration of the imconformity below the "Arecibo" 
is given on page 16, figure 3, of Berkey's paper. 
Berkey says in his summary of the geologic history of the island:' 

Where more simple marine conditions came into control, as would happen when 
submergence or planation had masked or destroyed the more elevated source of sup- 
ply, the deposits became almost wholly reef limestones and shell limestones, with 
only minor amounts of strictly detrital material irregularly distributed. 

The middle Oligocene reef-coral development of Porto Rico, there- 
fore, took place after its basement had been subaerially eroded and 
then depressed below sea level, and it seems that the basement prior 
to its submergence had been almost reduced to a peneplain surface. 

CUBA. 

Reef corals of middle Oligocene age were first collected in Cuba, 
on Rio Canapu, by Arthur C. Spencer, who obtained three species, 
all of which also occur in Antigua; but the only at all extensive col- 
lection is from the vicinity of Guantanamo, and was made by O. E. 
Meinzer, who studied in detail the stratigraphic relations of the coral- 
liferous formation. I am taking the following note from a manu- 
script by Mr. Meinzer, now awaiting publication. That there is a 
pronounced unconformity is indicated by a conglomerate at the base 
of the formation. Previous to the submergence, during which the 
coral reefs were formed, there was a long period of subaerial erosion, 
but geologic investigations have not been prosecuted over large 

1 Berkey, C. P., Geological reconnaissance of Porto Rico, New York Acad. Sd. Ann., vol. 36, pp. 12-17, 
1915. 

* Idem. p. 3. 

* Idem. p. 00. 



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262 



BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



enough areas in Cuba to draw inferences as to the physiographic 
features of the land surface resulting from the erosional activities. 

WEST INDIAN AND PANAMANIAN UPPER OLIGOCENE REEFS. 

ANOUILLA. 

Basic igneous rock above which in places there is some sandstone 
is exposed below the coralliferous limestone at Crocus Bay and 
Road Bay. The contact is very clearly one of erosion unconformity. 
The following is a composite of the sections exposed at Crocus Bay: 



Oeologie section at Crocus Bay^ Anguilla. 



3. Hard cavernous limeetone, with few or no corals 60 feet. 

2. More or less aigillaceoiis limestone with some beds of harder, purer lime- 
stone ; contains fossil corals from bottom to top, some coral heads as much 
as 2 feet in diameter; this member subdivisible into subordinate beds 
about 200 feet. 

1. Yellowish and brownish day underlain by dark blue-black clay, or sand- 
stone and conglomerate of igneous material overlying basic igneous rock 
(exposed at Pelican Point) 5feet±. 

The exposure at Road Bay is essentially the same as that at Crocus 
Bay. 

The AnguiUan reef was evidently formed during submergence after 
the subaerial erosion of its basement. 

It should be emphasized that the richly coralliferous limestone is 
overlain by more massive, harder, limestone in which there are few 
or no corals; and that the areal extent of the shoal-water Umestone 
indicates a submarine flat. 

CANAL 20NB. 

The Emperador limestone, according to Doctor MacDonald, lies 
unconformably on several of the beds belonging to the imderlying 
Culebra formation, and supplies another instance of a fossil coral reef 
with an imconformable basal contact. 

The stratigraphic relations of the important West Indian and Canal 
Zone reef corals and coral reefs are summarized in the following 
table: 

Stratigrapkic relations of West Indian and Canal Zone Eocene and Oligocene reef corals 

and coral reefs. 



Age. 


LocaUty. 


Basal contact. 


Overlying rock. 


Surfbce of base 
ment. 




perador Is.) 
Anguilla 


Unconformable on 
Culebra f ormatian. 

Unconformable on 
igneous rock or on 
sandstone and con- 
glomerate. 
do 






Upper OUgooene.... 


limestone without 
or with few corals. 

do 




Antigua..^ . . . ^ 


Da 




Porto Rioo(Peptno 

formation). 
Caba (Ouantanamo) 


... .do 


do 


Do. 


IClddle OUgooezie. . . . 


do 




Not known. 


Upper Eooene 


<»• 


Limestone without 
or withfew corals. 


Sutoierged flat. 



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GEOLOGY AND PALEONTOLOGY OF THE OANAL ZONE. 268 

AU of the fossil reefs discussed in the foregoing remarks were 
formed during periods of subsidence that followed subaerial erosion 
of their basements. The basal contacts might be interpreted as 
supporting Darwin's hypothesis, but in four of the six instances the 
reefs are buried imder later nearly pure limestones in which there are 
few or no corals. What caused the change in the character of the 
sediments, and coincidently led to the extermination of the reefs is 
not known; but the organisms in the overlying sediments indicate 
shallow, tropical waters, and as the geologic formations are areally 
extensive (relatively speaking), they were evidently formed on 
submarine flats. The corals b^an to grow on such flats and were 
ultimately killed. So long as the ecologic conditions were favorable, 
the corals flourished, but died when the conditions changed. The 
formation of the flats can scarcely be attributed to the corals. 

WEST INDIAN MIOCENE REEF CORALS. 

Meager developments of reef corals during the Miocene occur in 
Cuba and Santo Domingo, but at present no Miocene reefs are known 
unless the name reef be applied to the corals foimd in the La Ouz 
marl) eastward from La Cruz to the intersection of the railroad with 
the highway from Santiago to the Morro. The La Cruz marl is a 
bedded formation in which there are a few reef corals. The presence 
of pebbles in the basal part of the formation at the south end of 
Santiago Harbor suggests an erosion imconformity with some older 
Tertiary formation. 

No Pliocene reef corals are at present known in the West Indies. 
The erroneous suggestion, that a coralliferous limestone exposed in 
a quarry on Calle Infanta, opposite Castillo de la Pimta, Habana, 
might be PUocene, has been corrected on page 224. This limestone 
seems to represent yery nearly the same horizon in the Miocene 
as the Bowden marl of Jamaica; it may be stratigraphically somewhat 
higher. It contains some corals of reef f acies but it can not appro- 
priately be called coral-reef rock. The stratigraphic relations of the 
base of the deposit are not known. 

WEST INDIAN PLEISTOCENE REEFS. 

The West Indian Pleistocene reefs, whose stratigraphic relations 
have been critically investigated and can be discussed here are those 
of Jamaica and Cuba. Mr. R. T. Hill has placed in my hands a 
manuscript describing the Pleistocene reefs of Barbados, and Doctor 
MacDonald will discuss those of Costa Rica and Panama in his memoir 
on the geology of the Canal Zone and adjacent areas. 

The basal contacts of the Jamaican Pleistocene reefs, as has been 
elaborately presented by R. T. Hill in his account of the Jamaican ^ 

> Hin, R. T., The geology end physicAl geography of Jamaica, lios. Comp. Zool. Boll., vol. 84, pp. 90-09^ 
18Q9L 

37149—1^— Bull 108 6 



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264 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

reefs,, at least usually show unconfonnable relations. Although 
that Agassiz was aware of the imconf ormity at the base of the Cuban 
Pleistocene reefs can be inferred from his descriptions, he did not 
emphasize the stratigraphic relations; however, he does say r^arding 
the Uving Cuban reefs: ''In Cuba they [the coral reefs] abut upon the 
Tertiary limestone of its shores." I observed the imconformable 
relations at Baracoa, and stated that ''Upper Oligocene yeflowish 
calcareous marls or limestone are found in the vicinity of Nuevitas: 
also at Baracoa, where they immediately underlie the Pleistocene coasUd 
sohorruco."^ On page 32 of the same report it is stated: "It should 
be added here that all of the elevated Pleistocene coral reefs as seen 
by us and all of those recorded by those whom we consider compe- 
tent observers, are plastered on the surface of the upper Oligocene 
[mostly Miocene] formations, or in some instances upon older geolo- 
gic formations." 

Unconformable relations between the elevated Pleistocene reefs 
and the xmderlying Miocene limestone or marl are observable at 
Matanzas, Habana, and Santiago. The rock in the left for^round 
(pi. 71, fig. A,) is the slightly elevated soborruco (coral-reef rock) 
that extends into the mouth of Santiago Harbor, clearly show- 
ing that the harbor was outlined as a drainage basin previous 
to the formation of the particular reef now under consideration. 
The bluff and slopes in the background and on the right side of the 
illustration are formed in the Santa Cruz marl. 

The known un^^onformable relation at the base of the Pleistocene 
elevated reefs was the basis of inferred "subsidence of 80 to 100 feet" 
during the Pleistocene; this subsidence was followed by elevation and 
channeling in the mouth of the harbor; and this was followed by 
Recent submergence.* I have recently prepared a revised accoimt of 
the shore-line phenomena of Cuba, and present the following svmi- 
mary for the vicinity of Habana: 

1. Stand of land high enough for the subaerial erosion of the 
basement of a reef that seems to be about 30 feet above sea level 
at present, and for the outlining by erosion of Habana Harbor. 

2. Submergence in Pleistocene time to a stand about 30 feet lower 
than at present. 

3. Emergence in Pleistocene time sufficient to permit the cutting 
of a channel, now submerged 100 feet in Habana Harbor; the amount 
of this emergence would be about 100+ feet = 130 feet. 

4. Submergence, assigned to Recent time, to a depth of about 
100 feet. 

1 Hayes, C. W., Vaughan, T. W., and Spencer, A. C, A geological reconnaissance of Cuba, made under 
the dlrocti(m of General Leonard Wood, Military Governor, p. 23. The upper Oligocene in this quotation 
Is now considered Miocene. The italicised part of the sentence is in Roman letters in the original. 

*Idem., p. 34. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 265 

5. There may have been minor oscillations, for instance the 5-foot 
soborruco may represent slight elevation subsequent to the last 
submergence. 

Mr. O. E. Meinzer, in his manuscript, ''Geologic reconnaissance of 
a region adjacent to Guantanamo, Cuba," referred to on page 204, 
gives the following summary of events for the vicinity of Guantanamo : 

1. (Previous to the formation of the terraces) ''Erosion, resulting 
in the excavation of the principal valleys now in existence, some of 
them probably below present sea level. 

2. Submergence sufficient in amoimt to bring the land at least 
750 feet below the level of the present shore hue. 

3. Successive stages of emergence and perhaps slight tilting of the 
land, alternating with stages of quiescene, the emergence being about 
850 feet in. amount so that the land area stood about 100 feet higher 

> than at present, thereby permitting stream erosion below the present 
sea level; during the stages of quiescence sea benches and cliffs 
were formed at different, successive stands of the land. 

4. Submergence to the present level, resulting in the drowning of 
the lower parts of the stream valleys and in the production of innimi- 
erable small estuaries, bays, and coves. 

5. Filling of the submerged valleys and development of a new sea 
bench by destructive and constructive processes." 

The reefs considered in this section are fringing reefs. They rest 
unconformably upon their basements, but were formed during 
pauses in emergence. 

TERTL\RYAND PLEISTOCENE REEF CORALS AND CORAL REEFS OF THE UNITED 

STATES. 

SOUTHEASTERX UNITED STATES. 

In the United States Tertiary reef corals first appear at the base 
of the Eocene in the Midway group in Alabama, but these are not 
sufficiently abundant to entitle the deposit to the designation 
"coral reef." 

The oldest Tertiary coral reefs in this province are of middle 
OUgocene age, and have been studied at Salt Moimtain, near Jackson, 
Alabama, and near Bainbridge, Geoi^a. The basal contact of the 
reef at Salt Mountain is not exposed, and its nature is, therefore, 
unknown. The reef in the basal part of the Chattahoochee formation 
at Bainbridge, Geoi^a, rests on the surface of the upper Eocene Ocala 
limestone, which shows evidence of subaerial erosion, and is exposed 
from place to place along Flint' River throughout a distance of 8 
or 9 miles. It is relatively thin, perhaps only 10 to 15 feet thick, 
and contains a fauna of about 30 species of corals, mingled with 
which are many specimens of lAihotJiammon and large Lepidocydina. 

The next younger development of reef corals is in the upper part 
of the Chattahoochee formation and its stratigraphic equivalent. 



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266 BULLETIK 103, UNITED STATES NATIONAL MUSEUM. 

the ^'silex" bed and limestone of the Tampa fonnation. Corals are 
sufficiently abmidant to justify being designated ''reefs" at sevenJ 
localities, the most important of which are 18 miles south of Talla- 
hassee; Florida, in several counties in southern Georgia, and at 
Tampa, Florida. Ck>ralliferous limestone of the same or nearly the 
same age is exposed one-half mile south of River Junction, Florida, 
and at old Jacksonboro, Georgia. Well borings in Tampa show that 
beneath the coralliferous limestone is a variable thickness of clay 
which overlies the irregular surface of the Ocala limestone, indicating 
subaerial erosion, followed by submergence. The coralliferous beds 
are stratigraphically below the next younger set of deposits grouped 
under the Alum Bluff formation, indicating the continuation of ^ib- 
sidence after the formation of the reefs. The thickness of the reels 
and coralliferous beds is not great, perhaps between 10 and 20 feet. 
The f aima comprises about 20 species of corals. Where not siUcified 
and its character may be studied, the limestone associated with the 
corals is of complex origin. It is partly organic, probably in part a 
chemical precipitate, and contains terrigenous impurities. This indi- 
cates that the reefs and corals of this period grow during subsidence 
on a previously formed platform, but possess greater value for thrir 
aid in stratigraphic correlation than as constructional agents. 

The Alum Bluff formation, which, in my opinion, is of Miocene 
age, according to the usage adopted by the United States Greological 
Survey is subdivided into three members, which named from the 
bottom upward are the Chipola marl, Oak Grove sand, and Shoal 
River marl. The basal Chipola marl member was known only in 
an area extending from Aliun Bluff on Apalachicola River westward 
to Chipola River imtil it was recently identified by Miss Julia Gard- 
ner from a collection made by Dr. E. H. Sellards at Boynton Landing 
on Choctawhatchee River, in Washington Coimty. The bed on 
Chipola River seems conformably to overlie the Chattahoochee forma- 
tion, it is conformably overlain by higher beds of the typical Alum 
Bluff formation, and is between 15 and 17 feet thick. Of the four 
or five species of corals foimd at this horizon, one is of reef iacies, a 
massive species of Gomopora. Subsidence was in progress while 
these coralliferous beds were being deposited. 

Before completing the discussion of the Alum Bluff formation cer- 
tain events antecedent to its deposition in central peninsular Florida 
should be stated. Previous to the deposition of Chattahoochee and 
Tampa sediments, the Ocala limestone was deformed with the pro- 
duction of a low, elongate dome, the axis of which extends from 
near Gainesville to near Ocala. On both the east coast and the 
west coast along an east-west line through Gainesville the surface 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 267 

of the Ocala is below sea level and is overlain by younger forma- 
tions, while along the axis of the dome its surface rises from 80 to 
a little more than 100 feet above sea level. This low dome formed 
in the upper OUogocene sea an island or a group of islands to which 
I have applied the name ''Orange Island." The Chattahoochee 
and Tampa formations were deposited on the western slope of this 
island but they are not known in central Florida. The subsidence 
which brought about the deposition of these two formations con- 
tinued until the Alum Bluff sea advanced entirely across central 
Florida, where deposits of Alum Bluff age rest on the surface of the 
Ocala limestone apparently without the intervention of deposits of 
intermediate age. 

The portion of the Aliun Bluff formation above its basal member 
contains in central Florida at numerous locaUties heads of corals of 
reef facies belonging to the genus Siderastrea. At a place near 
Nigger Sink, about 8 miles north of Alachua, Florida, there is a 
Siderastrea reef, which, according to aneroid barometer measurement, 
is about 35 feet thick. The sediments associated with the Alum 
Bluff reef corals are greenish, usually phosphatic sands and clays, and 
impure phosphatic, in places magnesian limestone. The corals are 
decidedly subordinate in importance to other constructional agents, 
although they grew on a subsiding basement. 

Ahim Bluff sedimentation was succeeded by uplift and subaerial 
erosion preceding the depression initiating the deposition of the 
Choctawhatchee Miocene. Although the Miocene Choctawhatchee 
and Chesapeake faunas comprise about a dozen species of coials of 
distinctive facies, no reef corals are known as the temperature of the 
water was evidwitly too low. 

No Pliocene coral reefs are known, but corals of reef facies are well 
represented in the Caloosahatchee marl, which is largely composed of 
moUuscan shells. The stratigraphic relation of the Caloosahatchee 
marl to the Miocene has not been definitely ascertained, but available 
evidence suggests separation by an erosion unconformity. Whatever 
this relation may be, the formation was deposited during subsidence. 
Corals are of slight importance as contributors of material to the 
formation, as Heilprin long ago pointed out. 

The following table, which is a slightly revised copy of a table 
previously published,^ shows the stratigraphic distribution of coral 
reefs and reef corals from Oligocene to Recent time, and their rela- 
tion to changmg sea level. 

» Vsughan, T. W., and Shaw, E. W., Geologic investigations of the Florida coral reef tract, Carnegie 
Jntt, Waahingum Yearbook No. 14, p. 238, 1016. 



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268 



BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



Stratigrapkic diatrU>tUion of coral reefs and reef corals in the souiheaslem United States 
from Oligocene to Recent time, and their relation to changing sea level. 



Series. 



Becent 

pleistocene. 
Fliooene.... 



Miocene. 



Oligocene.. 
Eocene 



Geologic formations, members, 
unconformities. 



Distribution of reef corals and 
coral reefs. 



Oh OTgt* iff 

relatioiiof 

basement to 

sealev«L 



Erosion unconformity. 
Kev Largo limestone 

Erosion unconformity. 
Caloosah atchee marl 

Erosion unconformity. 
Choctawhatchee mari 

Erosion unconformity. 

Shoal River mari. 



Alum Bluff for-^ 
matioo. 



Oak Orove sand 
Chipolamarl.. 



Chattahoochee formaUon/LjSJ ; 



Erosion unconformity. 
Ocala limestone 



Coralreefs | Submergeooe. 

Coral reefs i Subsidence. 

Reef corals I Subsidence. 



No reefs, a few corals 

iA few corals; slight development of 
reefs in central and northern 
peninsular Florida. 
/A few corals; one species of reef 
\ faoies. 

Coral reefs (Tampa, Fla., etc.) 

Coral reefs (Bainbiidge, Ga.) 

No coral reefs 



Subsidence. 

Uubsidence. 

Subsidenoe. 
Subsideooe. 

Sul 



The table shows, besides the stratigraphic distribution of the reefs 
and reef corals, that, with possibly one exception, each development 
occurred during subsidence which followed subaerial erosion. 

To consider the basement of these fossil reefs: The geographic 
extent and composition of the limestones of upper Ek>cene age, which 
form the basement of the Floridian plateau, have been ascertained 
with considerable exactness. The surface outcrop has been mapped 
in Georgia and Florida, and well borings have revealed the presence 
of limestone of this age and character under yoimger formations in 
west Florida, at Panama City, and in Peninsular Florida, at Tampa, 
Key West, Key Vaca, and Palm Beach. The limestone is largely 
composed of the remains of Foraminifera, including myriads of 
Nummulites and orbitoidal Foraminifera, Bryozoa, and some mollusks 
and echinoids, with which is an imdetermined proportion of chemi- 
cally precipitated calcium carbonate and some terrigenous material. 
Corals are always rare and are usually absent. The organisms occur- 
ring in the formation are characteristic of tropical, shoal water, 50 
fathoms or less in depth. As the 100-fathom curve delimits the sub- 
merged border of the Coastal Plain, it is evident that the Floridian 
plateau was a part of the Coastal Plain and had essentially its present 
outline back in upper Eocene time before the formation of the oldest 
Chattahoochee reef, which was therefore superposed on a subsiding 
platform not produced by corals. The paleogeographic development 
of the Floridian plateau shows that each successive development of 
Tertiary reefs was on an antecedent platform which was formed by 
agencies other than those dependent on the presence of coral reefs. 
In all instances the volume of coral as compared with material from 
other sources is of minor and usually of negUgible importance. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 



269 



The accompanying map (fig. 7) shows the location of the Oligocene 
reefs with reference to the Plateau surface. 

That Pliocene deposition was followed by uplift, erosion, and de- 
pression, is shown by the fact that the Pleistocene shell marls along 
Caloosahatchee River rest on the eroded surface of the Pliocene. The 
Pleistocene reefs, the location of which is shown on the map (fig. 8) , 
were formed during subsidence which followed uplift at the close of 
Pliocene deposition. At the base of the reef, which is 105 feet thick. 




f 10. 7.— Florida, Ocala limestone plateau with suferposed Ouoocene and IIiocene coral reefs 

AND RK£r CORALS. Oc. 2«.=s0CALA LIMESTONE; THE FIGURES ARE FOR THE DEPTHS OF ITS UPPER SURFACE 
BELOW SEA LEVEL. C%=CHATTAH00CHEE AND TAMPA OUGOCENE FORMATIONS. A I. B.=ALVM BLUFF 

Miocene formation. 

is a calcareous deposit, 55 feet thick, of undetermined age. Beneath 
it are 450 feet of sand, mostly quartz, of Miocene age, below which 
foDow in descending order, limestones of Chattahoochee and Ocala 
age, but without any development of reef-corals. Planimeter meas- 
xu^ments indicate an area of 66 square mUes for the Pleistocene reef 
against an area of 1,670 for the chemically precipitated calcium car- 
bonate of the Miami and Key West oolites. I have already published 



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270 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

the statement that in Pleistocene time the calcimn carbonate chemi- 
cally precipitated probably predominated over that secreted by corals 
in the ratio 100:1.* • 

The theory advanced by Louis Agassiz * for the building of penin- 
sular Florida is familiar to most geologists through the writings of 
LeConte.' Agassiz says: ♦ ♦ ♦ ''the peninsula itself has once 
been a reef at least as far as the 28th degree of north latitude, as is 
shown by the investigation of the Everglades, and by the examination 




Flo. 8. FLOBIDA, location Or'FLEISTOCBNE CORAL KBXK A8 SHOWN BT +, AND THE LOCATION OF TEX 
AOASaB-LBCONTXBOXTNDABT OF 8VFP08BD OOBAL FOBKAIION. 

of the rocks at San Augustine." According to LeConte 's map about 
half of peninsular Florida was formed through the agency of coral 
reefs.* (See figure 8, above.) 

Eugene A. Smith, in 1881, showed that Eocene deposits extend 
south of Ocala into the peninsula; Heilprin showed that corals are 
imimportant to the latitude of Lake Okechobee; Alexander Agassiz 

> Vaaghan, T. W., Qlntdi of the geologio history of the Florida coral-reef tract and oomparisoii with other 
coral-feef areas, Joam. Washington Acad. 8ci, vol. 4, p. X, 1914. 
» U. S. Coast and Oeodetio Sunwy Ann. Bept. 1851, pp. 14&-160, 1852. 
* Elements of geology, ed. 4, p. 163. 



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OEOLOOY AND PALEOKTOLOQY OF THE CANAL ZONE. 271 

aooepted the results of Smith and Heilprin but contended that the 
sonl^em end of the peninsula is composed of wind-blown coral sand. 
Later investigations have established that the material comprising 
this part of the peninsula is neither coral sand nor is it wind-blown. 
Antecedent to the Recent reef out of an area of between 25,000 and 
30,000 square miles, perhaps as much as, but probably less than, 66 
square miles may now be attributed to coral. 

The data on the fossil reefs of the Southeastern States may be 
summarized as follows: 

1. Corals have played a subordinate part, usually a negligible part, 
in the building of the Floridian plateau. 

2. Every conspicuous development of fossil coral reefs or reef 
corals took place during subsidence. 

3. In every instance the coral reefs or reef corals have developed 
on platform basements which owe their origin to geologic agencies 
other than those dependent on the presence of corals. 

PUOCBNX BUT COBAL8 VSOM CABBISO CBKXK, CAIiTOBMIA. 

Mendenhall ^ has described in detail the relations of the coralli- 
f erous beds at this locality, and I have republished his statements 
in my account of the collection of corals made by him and Dr. Stephen 
Bowers.* There is here another instance of a richly coralliferous 
formation with an erosion imconformity at its base. 

LlTINO COBAL RbBN OF THB WbST InDIBS, FlOBIBA, AND ObNTBAL AmBBIOA. 

No general account of the position and general features of the 
Jiving reefs within the region above mentioned will be given here, as 
the subject has been fairly well treated by Alexander Agassiz for the 
West Indies and Central America,' and during the past eight years 
I have published a number of papers, listed in the footnote,^ on the 

1 Ifendenball, W. C, Notes on the geology d Oatrixo Mountain and Tidnlty, San Diego County, Gall- 
tenia, Joom. Oeology, yoL 18, pp. 836-365, 1910. 

t Vaoghan, T. W., The reef-oGral fiuina of Carrizo Creek, Imperial County, California, and its slgnifloanoe, 
» U. 8. OeoL Survey Prof. Paper 98-T, pp. 866-386, lAtes 0»-103, 1917. 

• Agassis, A., A reconnaissanoe of the Bahamas and of the elevated reefs ol Cuba in the steam yacht TF<tf 
Duck, Mus. Comp. ZooL BulL, vol. 96, pp. 146-166, 1894. 

« Vaughan, T. W.: 

Sketch d the geologic history of the Floridian Plateau, Sdenoe, new ser., vol. 82, pp. 24-37, July 1, 1910. 

A contribution to the geologic history of the Floridian Plateau, Carnegie Inst. Washington Pub. No. 133, 
pp. 99-186, 1910. * 

Studies d the geology and of the Madreporaria Of the Bahamas and of southern Florida, Gamegie Inst. 
WasUngton Year BocA No. 11 (for 1912), pp. 168-162, 1913. 

Bemarks on the geology of the Bahamas and on the formation of the Floridian and Bahaman oolites, 
WasUngton Acad. SoL Joum., vol. 3, pp. 802-304, May 19, 1913. 

With L. V. Pirsson, A deep boring in Bermuda Island, Amer. Jour. ScL, ser. 4, vol. 36, pp. 70-71, July, 
1013. 

Sketcfaofgeologichistoryofthe Floridaooralreeftractand comparisons with otherooralreef areas, Wash- 
ingtoo Acad. SoL Joum., voL 4, pp. 26-34, Jan. 19, 1914; abstract, Oed. Sec. America Bull., vol. 26, pp. 
41-42, March, 1914. 

The reef corals of southern Florida, Gamegie Inst. Washington Year Book No. 12 (for 1913), pp. 181-188, 

1914. 

(Footnote continued on page 272.) 



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272 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Floridian, Bahamian, West Indian, and Central American reefs. 
In addition to my studies in the field and my work on charts and 
maps in the office, I have compiled all. available information on 
Pleistocene and Recent strand-line movement along the Atlantic 
coast between Argentina on the south and New England on the 
north. 

The discussion to follow will present evidence on Recent change in 
the position of strand line, on the amount of change, and on the rela- 
tions of the living coral to the basements on which they have formed 
for the West Indies from Antigua along the Caribbean arc to Cuba, 
the Bahamas, the Bermudas, Florida, and Central America. Accounts 
of these areas will be followed by remarks on some other West Indian 
Islands, on the Brazilian reefs, on the Argentine shore line, and on the 
shore line of the United States between Florida and Cape Cod. 

[Footnote contiiitiod Crom pAfi 271.) 

Investigations of the geology and geologic proooesses of the reef tracts and adjacent areas in the BalmmOT 
and Florida, Carnegie Inst. Washlngtoo Year Book No. 12 (for 1013), pp. 188-184, 1914. 

The platforms of barrier ooral reefs, Amer. Goog. Soc. BolL, voL 46, pp. 42^-429, 1014. 

Preliminary remarks on the geology of the Bahamas with special relbrenoe to the origin of the Fktridian 
vid Bahama oolites, Carnegie Inst Washington Pab. No. 182, pp. 47-54, 1014. 

The building of the ICarquesas and Tortogas atoUs and a sketch of the geok)gic history of the Florida 
ree; tract, Carnegie Inst. Washington Pab. No. 182, pp. 5&-^, 1914. 

Study of the stratigraphio geology and of the fbssil corals and associated organisms in several of the smaller 
West Indian Islands, Carnegie Inst. Washington Year Book No. 13 (tot 1014), pp. 356-360, 1915. 

Qeologioal investigations in the Bahamas and southern Florida, Carnegie Inst. Washington Year Book 
No. 13 (for 1014), pp. 227-233, 1015. 

Reef corals of the Bahamas and soathem Florida, Carnegie Inst. Washington Year Book No. 13 (fbr 
1914), pp. 222-226, 1015. 

Coral reef^ and reef corals of the southeastern United States, their geologic history and their signlflcanoe. 
Abstract, Science, new ser., vol. 41, pp. 506-500, April 2, 1015; Oeol. Soc America BuL, vol. 26, pp. 58-60^ 
1015. 

Introductory remarks to symposium on the fiactors producing changes in position of strand line during 
the Pleistocene and post-Pleistocene, Washington Acad. Sci. Joum., vol. 5, pp. 444-445, June 18, 1015. 

[R^sum6 of the present status of the geologic correlation of the Cretaceous and Tertiary formati<Mis of the 
AntUles], Washington Acad. ScL Joum., voL 5, p. 480, July 10, 1015. 

Memorandum on the geology of the ground waters of the Island of Antigua, B. W. I., West Indian BulL, 
vol. 14, No. 4, 4} pp., 1015. Imperial Dept. of Agri. for the West Indies. 

The geologicsigniflcance of the growth rate of the Floridian and Bahaman shoal-water corals, Washington 
Acad. Sci. Joum., vol. 5, No. 17, pp. 501-600, Oct. 10, 1015. 

On Recent ICadreporaria of Florida, the Bahamas, ami the West Indies, and on collections from Murray 
Island, Australia, Carnegie Inst. Washington Year Book No. 14 (for 1015), pp. 220-231, 1016. 

And Shaw, E. W., Geologic investigations of the Florida coral-reef tract, Carnegie Inst. Washington 
Year Book No. 14 (for 1015), pp. 233-238, 1016. 

Study of the stratigraphic geology and of the fossil corals and associated organisms in several of the smaller 
West Indian Islands, Camegie Inst. Washington Year Book No. 14 (for 1015), pp. 368-373, 1016. 

Present status of the investigations of the origin of barrier coral reef^ Amer. Joum. Sci., ser. 4« voL 41, 
No. 241, pp. 131-135, January, 1916. 

The results of investigations of the ecology of the Floridian and Bahaman shoal-water corals, Nat. Acad. 
Sci. Proc., vol. 2, pp. 05-100, February, 1016. 

Some littoral and sublittoral physiographic features of the Virgin and northem Leeward Islands and 
their bearing on the coral reef problem, Washington Acad. Sci. Joum., vol. 6, No. 3, pp. 53-66, Feb. 4, 1016i 
also abstract Geol. Soc. America Bull., vol. 27, No. 1, pp. 41-45, 1016. 

The corals and coral reefs of the Gulf of Mexico and the Caribbean Sea (abstract of paper read before 
special meeting of Amor. Ass. Adv. Sci., in cooperation with Pan-American Congress), Science, new ser., 
vol. 43, pp. 250-251, February 18, 1016. 

In collaboration with Cnshman, J. A., Goldman, M. I., Howe, M. A., and others: Some shoal-water 
bottom samples fh>m Murray Island, Australia, and comparisons of them with samples from Florida and 
the Bahamas, Camegie Inst. Washington Pub. No. 213, pp. 235-207, pis. 04-08, 1018. 

Chemical and organic deposits of the sea, Oeol. Soc American Bull., voL 28, pp. 933-944, pis. 47, 4S, 1918. 



Digitized by VjOOQIC 



GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 273 

ANTIGUA-BARBUDA BANK. 

The islands of Antigua and Barbuda rise from a bank which is 
bounded by the 100-fathom curve and is 50 miles in length along a 
north-south line, and from 13 to 20 miles in width. Antigua is near 
the southern and Barbuda near the northern end, with water from 
15 to 18 fathoms in depth between them. (See text fig. 9.) 



'^:.' 

'*§: 

r 
'« 



133 n» 



38 
15 

13 
18 

18 i: 

12 
35 



i Xuioo.Rk' 
6/4 



^4K 



lot' 



•11 



225/30 19 17 ^g 



, NBARBUDA. 

lb\ (British 



13 i 

PaU. 



ipunUfaJH. 



aUaettoJ'Cf^3^^*CT'h8 f 
13 ^** W .i3 liji^ ' »J' 33?^ 

^2 24 ,5 jh/^-m / 

15 M C».Sfc.^Vrtfl>-'lj5 ia/l« 



"^^8 



19 
18 
Ifi \3j20 
14 16 16 1 



15 






15 15 



18 



17 
15 > 17 



U \ 4 

15 21 \ 

17-^V^l^ »»• 








16- , 

''1 ANTIGUA 

T/lo (Brirish) 






3t^^^^^.^^ 



17' 



Fig. 9.— Cuart op axtiova— barbuda bank. From U. S.'htdrooraphic chart no. 2318. Scale,! 
inch- about 12.8 nautical miles. 

The shore line of Antigua is deeply indented by numerous bays 
and harbors, as St. John, Five Islands, and Falmouth harbors, and 
Willoughby, Nonsuch, and Belfast bays (pi. 68, figs. A, B, and 
text fig. 10). The absence of terraces and elevated wave-cut cliflFs 
is especially noteworthy. The discovery m St. John Harbor, at a 
depth of .20 feet below sea level, of a 4-foot bed of peat, which 



Digitized by VjOOQIC 



274 



BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



is not composed of marine plants (according to Mr. C. A. Davis), 
adds confirmation to the inference from the indented shore Une and 
the absence of elevated terraces and wave-cut cliffs that the last 
important movement of the strand Une was one of submergence. 
Present sea-level relations have persisted long enough for the develop- 
ment of sea cUffs, in places 100 feet or more high, for the alluvial 
fillings at the heads of the bays, and for the extension inland of 
alluvial deposits along the streamways. There is some evidence of a 
slight upward movement of the land, a few feet, less than 10, since 
the submergence. 




Fio 



10.— CHABT or PAET OF BAST COAST Of ANTIOUA. FROM U. S. BTDBOORAPHIC CHART NO. lOM. 



Barbuda, which is composed of limestone and has a maximum 
height of about 200 feet, has no marked tudentations of its shore 
line; but Dr. H. A. Tempany informs me that fresh-water springs 
emerge below sea level in the lagoon about one-half mile south of 
Codrington village, a fact of significance in probably indicating 
submergence. 

The similarity of the land mollusca of Antigua and Barbuda lend 
support to the inference from physiographic data that these islands 
were part of one land mass in Pleistocene time and have been severed 
by submergence, and as the water between the islands is 18 fathoms 
deep, the sea level must have risen at least that amount. A sub- 
merged steep slope off the southeast side of Antigua at depths be- 
tween 100 and 150 feet accords with submergence to a depth of at least 



Digitized by VjOOQIC 



GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 



275 



18 fathoms, and indicates submergence of about 120 feet or 20 
fathoms. (See fig. 11 below.) 

Barrier coral reefs occur around Antigua off the mouth of Nonsuch 
Bay, off the southwest angle of the island, and there is a discon- 
tinuous barrier off the west side of the island. There are other reef 
patches, some of which are almost barriers. Barbuda has barrier 
roefs, Cobb and Groat reefs, off its northern end. 

These reefs of Antigua and Barbuda ocoiu* on a platform which has 
been submerged. That the platform or flat lying between Antigua 
and surrounding Antigua is in origin independent of the corals growing 
on its surface is shown not alone by its continuity irrespective of 
the presence of corals. That a land area existed between Antigua 
and Barbuda in Pleistocene time is clearly shown by the land mol- 
lusca; while the submerged steep slope or scarp shows that the flat 




SOUTHEAST COAST OF ANTIGUA 



Ses /eye/ 




t eJ^mA ► 

NORTH COAST OF 


HAVANA HARBOR 


MOSOUITO BANK 


ST. THOMAS 


shovving depth of filled 
ct»nn«l In harbor 





FlO. 11.— SUBMABINE PBOFILXS OWT WEST INDIAN IBLANDS AKD ACROSS MOSQUITO BANK. 

existed and was marginally cut by the sea while it stood about 120 
feet higher than at present. 

ST. MARTIN PLATEAU. 

JL W. Spencer has applied this designation to the plateau on which 
St. Bartholomew^ St. Martin, and Anguilla stand. This plateau, as 
bounded by the 100-fathom curve, is irregular in shape and is 75 
miles long by 45 miles wide. The maximum depth of water between 
St. Bartholomew and St. Martin is 16 fathoms and between St. 
Martin and Anguilla 14 fathoms. (See text fig. 12.) 

The shore line of St. Bartholomew is indented, the indentations are 
usually divided by beaches into an inner or lagoon part and an outer 
bay or harbor part (pi. 68, figs. C, D). The beaches may have 
been elevated between 3 and 5 feet. The lagoons behind the beaches 



Digitized by VjOOQIC 



276 



BULLETIN 103, UNITED STATES . NATIONAL MUSEUM. 



are the salt ponds of the island. There is an entire absence of ele- 
vated terraces, unless some apparent shoulders on outlying islete, not 
actually visited by me, should be slightly elevated sea-cut benches. 
Wave-cut cliflfs margin the rocky shores, and alluvial flats occur 
around the heads of the bays. 

llie shore line of St. Martin is indented. Each reentrant into the 
land is usually divided by a transverse beach into an inner lagoon or 

flfl! 



il 



801 



M 



I Co. 986 »\ 
19 :»^ r4 



/ 



nag. 



|40\ 



- 27 40 N. 

31 34' 

wk.S. 



1^^86 



^r"^^- 



25 23 



37 »^38--^®^ 



13 _'>4 







tlG. 12.— ^liAKT U*' ST. UAAim VLATXAU. FBOM U. §. ETDSOOBAPHIC CHART HO. 2318. bCALK, OXB 
INCH— ABOUT 12.8 NAUTICAL ITILES. 

salt pond and an outer bay portion; and alluvial flats margin the 
heads of the reentrants and project inland between the hills. The 
spurs along the shore are truncated by wave-cut cliffs (see pi. 69, 
%. A) and exhibit no definite terracing. Older beach rook was seen 
at the northeast end of Blanche Point, perhaps indicating slight 
differential uplift for that locality. 

The shore line of AnguiUa (see pi. 69, figs. B, D), although not 
so conspicuously indented as that of St. Bartholomew and St. Martin, 



Digitized by VjOOQIC 



GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 277 

is indented, and a number of instances, Road Bay, for example, of 
the separation by beaches of an inner lagoon from an outer bay are 
present. Three instances of inclosed basins having underground 
conununication with the sea were noted (pi. 69, fig. C). No definite 
terraces are present and wave-cut cliffs are greatly developed. 

That the last important change of sea level was by submergence 
of the land is evident from the character of the shore line in St. Bar- 
tholomew, St. Martin, and AnguiUa; and in Anguilla additional 
evidence is afforded by the undergroimd communication between 
inclosed basins in the limestone and the sea. Stable condition of the 
shore line for a considerable time is attested by the wave-cut cliffs, 
the development of the beaches, the alluvial fillings at the heads of 
reentrants into the landmass, and in St. Martin by the presence of 
unterraoed flood plains along the streamways. 

In my paper on the littoral and sublittoral physiographic features 
of tile Virgin and northern Leeward Islands, referred to in the foot- 
note on page 272, 1 have shown that on the windward side of the St. 
Miyiin plateau there is an outer deeper flat, 26 to 36 fathoms below 
sea level, with a maximum length east and west of over 30 miles, and 
that this flat may be subdivisible into two subordinate terrace flats. 
The scarp on the landward side of the deeper flat in places is about 
50 feet hi^, in depths between 20 and 28 fathoms; above the deeper 
flat is a shallower one, whose outer edge is about 20 fathoms under 
the sea (see text-fig. 11, p. 276). Other submarine evidence of sub- 
mergence in this area is given in my paper cited. At the time the 
shore line around the St. Martin Plateau was about 20 fathoms 
lower than at present, Anguilla, St. Martin, and St. Bartholomew 
must have been united. The biologic evidence at present available 
is not sufficient to be decisive, but all that is known accords with 
this interpretation. Notches on the outer edge of the plateau sim- 
ulate hanging valleys and may represent the outer ends of valleys 
out while the sea stood about 40 fathoms lower than now; but the 
information on these is too scant to justify more than a suggestion. 

The hydrographic chart does not show well the reefs of these islands, 
nor does the British Admiralty West India Pilot give a good descrip- 
tion of them. Because of rough weather most of my own observa- 
tions were made from the shore. Coral reefs occur across the en- 
trances to most of the bays on the northeast and southeast sides of 
St. Bartholomew; reefs are well developed on the east side of St 
Martin, off North Point, and on the southeast side of Tintamarre 
Island; and there are dangerous reefs off the southeast coast of 
Anguilla and on the north coast of the east end of the island. Seal 
Island reefs occur on a ridge extending westward from the northeast 
end of Anguilla. Some of these reefs are of the barrier type, as 
navigable channels lie between them and the shore, one at Forest 
Point is an instance. 



Digitized by VjOOQIC 



278 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

The reefs of the St. Martin Plateau are superposed on an ante- 
cedent platform that was brought into its present relations to sea 
level by geologically Recent submergence to an amount of about 20 
fathoms. 



65' 



18 



1 olUrn frequently occur / 29 86 IT 12 4X*— ■»»%— 



» ao 



30 31 



\X%, -^-*il. 

^, — ''so 86 88 "-*, 



J^I^^^^^^T^^nwAD 



/4l 



:io Ji •^'' 19 



SI 



'» \; " a » 18 IS ^j 



« (r8)«s 






27 



30 

27" Si • m 















IIM 










I 

23 28 
..S3 2I_ «1 



<» 



980 ' 



no 



85 



-^29 



610 



326 



1105 



18» V 

V 13W 



^ ^ 1001 
1337 



876 

"^ 1350 

1U6 610 

»90 

006 



1698 

— ma- 



im 



1091 
8U 



981 






1870 



UO 



AGO 



9J6 ^ 21V _rr-^^ a;i 



ST-CROIX \ 

^ 2 F. r. 




' ll7""Tl2 



Fxo. 13.— Chart of tiboik islands akd st. cboix. From U. S. htdboobaphic chart no. 2318. SotSi 

ONS INCH^ABOUT 12.8 NAUTICAL MILES. 

ST. CROIX ISLAND. 

This island rises above a bank about 30 miles long and 10 nuks 
wide. The distance from the shore to the 100-f athom curve is usually 



Digitized by VjOOQIC 



GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 279 

less than three-quarters of a mile on the west end; and on the north 
side west of Sugar Bay the distance ranges from one-quarter to one- 
half mile. 0ff the south shore the distance to the 100-fathom curve 
in places slightly exceeds 3 miles; off the east end for 7 miles the 
water is less than 40 fathoms deep, while off the north coast the 
platform gradually narrows westward until near Salt River Point its 
width is less than one-half mile. 

There is a long, disconnected barrier reef off most of the south 
coast, and barrier reefs are present off the north coast to a short dis- 
tance west of Christiansted. The indented, ragged coast line and 
the depth of water on the platform so clearly point to the same con- 
clusion as that already drawn from a study of Antigua, St. Bar- 
tholomew, etc., that reiteration is not necessary. 

VIRGIN BANK. 

The Virgin group of islands consists of about 100 small islands and 
keys (text fig. 13). The bank above which they rise is an eastward 
prolongation of that on which Porto Rico stands. The chart shows the 
indented coast Ime and the extensive, relatively shoal platform above 
the surface of which the islands project. The maximum depth of 
water between the islands is about 17 fathoms. St. Thomas well 
exhibits the coastal phenomena to which attention has already been 
so often directed — ^reentrants with alluvial fillings at their heads, 
unterraced alluvial bottoms along streamways, and wave-cut cliffs on 
the imterraced promontories (pi. 70, figs. A, B, G). 

In my paper on some littoral and subUttoral physiographic features 
of the Virgin and northern Leeward Islands, already referred to, it 
has been pointed out that there are three terrace flats under the sea 
off St. Thomas, St. John, Tortola, and Virgin Gorda (see text fig. 11, 
p. 275). On the leeward side the deepest lies between 26 and 30 
fathoms in depth and is separated by a scarp or steep slope on its 
landward side from A flat ranging from 14 to 20 fathoms in depth, 
which in turn is separated by a steep slope from a flat ranging from 
6 to 10 fathoms in depth. On the windward side the respective 
depths are 26 to 34 for the deepest flat, 14 to 20 fathoms for the 
intermediate flat, and 7 to 10 fathoms for the shallowest one. The 
intermediate flat is narrow or absent on the promontory tips on the 
windward side, while it is preserved on the leeward side, strongly 
suggesting, if not actually proving, that the intermediate flat is older 
than the deeper one and was cut away in exposed places while the 
deeper one was forming. This evidence necessitates •the deduction 
that in recent geologic time the Virgin Islands, except minor differ- 
ential cnistal movement in the vicinity of Anegada, have been sub- 
merged to a depth of about 20 fathoms, and that they were previously 
joined to Porto Rico, a deduction completely corrobdrated by bio- 
87149— 19— BqU. 108 7 



Digitized by VjOOQIC 



280 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

logic evidence, for Dr. L. Stejneger says in his herpetology of Porto 
Eico that "St. Thomas and St. John form only a herpetological ap- 
pendix to Porto Rico/' and Dn P. Bartsch informs me that the 
testimony of the land moUnsca is the same as that of the reptiles 
and batrachians. Indentations at depths of abont 40 fathoms in the 
outer edge of the submarine bank simulate hanging valleys that may 
have been formed while the sea level was 40 fathoms lower than at 
present. 

In the Virgin Islands there are three tiers of coral reeis, namely, 
(1) on the outer edge of the deepest flat, (2) on the outer edge of the 
intermediate flat, (3) within depths of 10 fathoms or less. The ree& 
could not have been formed on the deepest flat while the scarp on 
the landward side of the flat was being cut, and the other reefs are 
clearly younger than the basements above which they rise, for their 
basements existed and had had a complicated history prior to 
the formation of the living reefs. In fact, the basements were dry- 
land surfaces during at least a part of Pleistocene time. 

CUBA. 

The principal contributors to the literature on the shore-line phe- 
nomena of Cuba are W. O. Crosby,* Alexander Agassiz,* R. T. Hill,* 
Vaughan and Spencer,* and Hayes, Vaughan, and Spencer.* I have 
in papers cited on pages 271, 272 referred to some of the features of 
the Cuban shore line as bearing on the conditions under which the liv^ 
ing coral reefs off the shores of the island have foimed. W. M. Davis 
has recently alluded to the origin of the pouch-shaped harbors,* and 
here it may be weU to direct attention to a criticism made by him 
in his article cited in the foot note. He says: 

It is, however, worth noting that the embayments here considered have a quite 
different relation to the adjacent coral reefs from that found, according to Hayes, 
Vaughan, and Spencer, in the pouched-reef ^ harbors of Cuba: All the embayments 
I saw inside of sea-level barrier reefs in the Pacific islands occupy valleys older than 
the reefs; but in Cuba the valleys, and still more the subsidence which drowned 
them in producing the pouched harbors, are described by the above-named authors 
as younger than the elevated reefs which inclose them; and such valleys do not bear 
on the origin of the reefs, as appears from the following extract: * * * 

The extract is followed by comment, then by a quotation from 
Crosby and one from Hill, after which he says: "Without additional 

1 Crosby, W. O., On the elevated reefs of Calm, Bost. Boo. Nat. Hist. Proc., vol. 22, pp. 134-130, 1883. 

< Agassic, A., A reoonnaissance of the Bahamas and of the elevated reefs of Cuba In the steam yacht 
wad Dvck, January to Apiil, 1803, Mos. Con^>. Zool. BoIL, vol. 26, pp. 10»-1M, 18M. 

s Hill, R. T., Notes/>n the geology of the island of Cuba, Mas. Comp. Zool. BuU.^ vol. 16, pp. 278-281f 
1895. 

* Vaughan, T. W., and Spencer, A. C, The geography of Cuba, Amer. Geog. Boo. Boll., voL 34, pp. 106- 
116, 1902. 

» Hayes, C. W., Vau^an, T. W., Spencer, A. C, Heport on a geological reoonnaissttice of Cuba, pp. 
123, 1902. 

• Davis, W. M., A «haler Memorial study of coral reef^, Amer. Joum. 8d., ser. 4, voL 40, pp. 227-428 
1915. 

T "Pouched-reef harbors" are words not used in the pubUcatiom under dlsoosstomby ProflBOor Davis. 



Digitized by VjOOQIC 



GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 281 

field study it is impossible to say which one of these views is correct, 
but the features of the Pacific reefs that I have seen support Hill's 
explanation." I have twice published the statement that ^' Hayes, 
Vaughan, and Spencer have shown, as is evidenced by the pouch- 
shaped harbors of the Cuban coast and filled channels, such as the 
submerged filled channel in Habana Harbor, that the last movement 
of the Cuban coast has been downward with reference to sea level," 
and that "the platform on which the Cuban reefs grow^ has been 
brought to its present position by subsidence." These remarks 
apply to the present Uving coral reefs and not to the elevated reefs, 
and the conditions presented by the pouch-shaped harbor is only a 
part of the evidence showing recent submergence of the Cuban shore 
line. 

Professor Davis's remark that " all the embayments I saw inside 
the sea-level barriers in the Pacific occupy valleys older than the 
reefs" has no appHcation whatever to the protecting effect a fringing 
reef may have on the shore of a land during elevation subsequent 
to the formation of a fringing reef, thereby permitting erosional 
agencies to operate more rapidly on the softer rocks lying back 
from the shore. The words in the Cuba report are: "Wherever the 
conditions are favorable for the growth of corals a fringing reef is 
built * * *." 

On preceding pages of this paper I have shown that there were 
coral reefs in Cuba in middle Oligocene time^ that there were reef 
corals in both upper Oligocene and Miocene time (this Miocene is 
called upper Ohgocene in the Cuba report); and that there are 
Pleistocene as well as living reefs. In the Miocene La Cruz marl in 
the vicinity of Santiago the greatest abundance of reef corals is not 
at the present head of Santiago Harbor, but it is seaward of the town 
of Santiago, east of La Cruz. (For a view seaward through the 
mouth of Santiago Harbor, see pi. 71, fig. B.) Whether the coral 
heads are sufficiently abimdant to have retarded erosion toward 
the mouth of the harbor, while it was more rapid on the landward 
side, I am not prepared to say. This, however, was not a fringing 
reef, should it be appropriately considered a reef. 

As to whether the elevated Pleistocene fringing reefs extended up 
to the sides of the outflowing water at the harbor mouths, thereby 
maintahiing restricted outlets, or whether channels have been cut 
across the reefs after uplift, either of the alternatives is possible. 
Off the mouths of bays in Antigua, channels are maintained across 
living barrier reefs, which are tied to the shore at one end; while 
off Virgin Gorda, a barrier reef extends perpendicularly across the 
axis of the mouth of a submerged valley. These are living reefs, 
which have grown up 'during or after submergence and are yoimger 

1 Not italidxed in tbe origixiaL Note use of present tense, " grow,** 



Digitized by VjOOQIC 



282 



BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



than the valleys landward of them. However, as the elevated 
Cuban reefs under consideration are fringing reefs, it seems to me 
more probable that they never extended across the harbor mouths; 
and I will add that the harbor basins had been formed, at least in 
large part, before the development of the now elevated fringing 
Pleistocene reefs. 

Crosby, in 1883, seems to have been the first one to recognize the 
significance of the pouch-shaped harbors of Cuba. He says:* 

* * * During this period of elevation, Cuba, like most rising lands, had few 
harbors, but when subsidence began the sea occupied the channels and basins which 
had been excavated and cleared out by the rivers, and thus a laige numb^ of harboza 
came into existence. * * * They are half-drowned valleys filled to a consider- 
able depth with land detritus, conditions which could not exist if the land was rising 
or had risen. 



There are very many pouch-shaped along the Cuban coast, 
following table presents information on 15 of them: 

Principal Cuban harbors. 



The 





Shape. 


Maxi- 
mum 
width, 

sea- 
miles.! 


Maxi- 
mum 
known 
depth 
in chan- 
nel or 
harbor. 


Chan- 
nel 
length, 
sea- 
miles. 


Chim. 
nel 


Hei|^t<tfadK. 
cent land. 


Name. 


narrow- 
est 
part. 


East 
side. 


West 
side. 


NOBTH COAST. 

Bahia Honda 


Palmately digitate 


3.00 
0.00 
1.60 
«2.00 
.7.00 
7.60 
3.26 


Feet 
«5» 

- 79 
172 
iOO 

•137 
75 
85 

»234 

"168 
166 

W33 
59 

68 

68 

W189 


1.60 

.60 

.60 

.76 

4.38 

1.76 

1.60 

•2.00 

.60 

.63 

.18 
3.76 

.38 


FeeL 

2,180 

1825 

900 

470 

1,400 

900 

460 

000 

800 
6,680 

676 


FmL 
80-40 
160 


FkeL 
60 


Cabafias 


Trilobate 


too 


Hariel 


Irregularly diidtate 




Havana . . 


TSS5ate...7r.....::::::..:: 


200 

Flat. 
Flat 
• 100 
•200 
50-75 
120-176 

500 
436 

230 


±10 


Nuflvltaa 


Bilobate 


^t 


Padre 


Irregularly bflobate 


TFUt. 


Banes 


Palmately digitate 


•150 


Nipe 


Unequally bifobate 


• 200 


Llvlsa and Cabonico. . . 
Tanamo . < . . . . 


do 

Irrogularly bilobate 


8.00 
5.88 

.00 
5.00 

1.00 


75-100 


SOUTH COAST. 

Baitiqaeri 


Trilobate head 


000 


Ouantanamo 


Irregularly dumb-bell 
TTtiilatwiU 


810 


Santiago 


220 


Bnseflada de Mom 






Cienfuegos 


TTniiAteral 


4.25 


2.13 


1,200 


180 


157 









1 1 sea-mile-6,081 feet. 

• 1 10 feet outside at channel mouth. 

• 90 feet in channel mouth. 
< About. 

• Submersed channel 100. 

• 100 feet frequent. 

' Coral rock according to A. Agassis. 



• South. 
•North. 

u 180 feet and over frequent. 
" 150 feet frequent. 
!• 78 feet at mouth, 
u 214 feet off Pta. Pasa CabaUot. 



It is important to note that where the harbors are digitate in 
shape, Bahia Honda for instance, one or more streams enter each 
digitation, and that the mouths of the streams are either embayed 
or, in places, swamps and delta plains have formed. The pouch- 
shaped harbors are not the only indentations of the shore line, for 
the lower courses of all the larger streams are more or less embayed. 

1 Crosby, W. O., On the elevated reefs of Cuba, Bost. Soo. Nat. Hist. Proc., voL 24, pp. 124-130^ 1883. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 



283 




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284 BULLETIN 108, UNITED STATES NATIONAL MUSEUM. 

How are the harbors to be explained? Doctor Hayes and I 
beUeved we found the answer in the conditions at present existing 
along Yumuri River, near Matanzas. The river here empties into 
the sea through a narrow gorge cut through Miocene limestone and 
marls (see pi. 71, fig. C). The top of the gorge is 200 feet above 
sea level, while farther back from the stream altitudes of 400 feet 
or sUghtly more are attained. Above the gorge, the Yumuri and 
its tributary, Rio Caico, have simk their courses through the Ume- 
stone, have removed it, and have developed wide, almost base-level 
valleys (see pi. 71, fig. D), on the imderlying softer sandstone 
and shale. If this basin were depressed sufficiently to let the sea 
into it through Yumuri gorge a pouch-shaped harbor would result. 

Additicmal evidence bearing on the problem of the origin of these 
harbors was obtained from records of borings. Mr. C. A. Kiiowlton, 
an engineer at Santiago, reported to us that in boring wells in the 
valley of San Juan River, 3 miles southeast of Santiago, he found 
at a depth of 70 feet below sea level what appeared to be stream 
gravel. Even more convincing evidence was obtained in Habana 
Harbor. In the preparation of plans for a sewerage system the 
Military Grovemor had a series of borings made across the harbor. 
This harbor occurs in a rather wide vaUey surroimded by sides 
which slope upward from sea level to an altitude of about 200 feet. 
The borings revealed a submerged terraced valley within the wider 
valley and in the middle of the inner valley a channel reaching a 
depth of more than 30 meters (about 100 feet) below sea level (see 
text-fig. 14). The depth of the first flat above the sides of the 
channel is about 13 meters (about 42 feet) below sea level. This flat 
is now covered with sand and the submerged channel is filled with 
sand and clay. There are at present no known processes whereby 
such a channel and terrace could be developed and then buried, 
except by a higher stand of the land enabling a stream to cut a 
trench and develop a terrace, followed by a lower stand of the land 
which submerged both the channel and the terrace and resulted in 
their burial by sediment deposited over them. 

It appears to me that there is no escape from the interpretation, 
made first by Crosby, that the pouch-shaped harbors are drowned 
drainage basins. Before the accimiulation of the data by Hay^, 
Spencer, and me. Hill endeavored to explain them without a shift 
in height of strand-line, but after the additional information was 
presented to him he abandoned his interpretation and accepted ours. 
There is a statement to this effect in a manuscript by him now in my 
possession, and this citation is made with his authority. 

The factors producing the peculiar form of the harbors will now 
be briefly considered. According to Crosby, Hill, and the account 
in our report on Cuba, fringing reefs are supposed to have restricted 



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GEOLOGY AlH) PALEONTOLOGY OF THE CANAL ZONE. 285 

the mouihs of the streams, either by growing up to the edges of the 
outflowing water, a channel thereby being maintained, or because of 
their greater hardness they offered greater resistance to erosion than 
did the softer rocks on their landward side. It is my present 
opinion that the hypothesis of the reefs having more than a secondary 
importance in the development of these features must be discarded 
for the following reasons: First, that such physiographic forms are 
in no wise dependent on the presence of coral reefs is shown by their 
frequency in areas imderlain by Cretaceous limestones in Texas. 
HiUcoat Valley in the southwest quarter of the Nueces quadrangle, 
Texas, is such a basin, with a narrow outlet into Nueces River. This 
is only one of a niunber of instances that might be given. ^In phy- 
siographic form this basin and its outlet resemble the pouch-shaped 
harbors of Cuba. Second, there is no evidence that corals had any 
more influence in Cuba than in Texas, for instance, Yiuniui gorge at 
Matanzas is about 200 feet deep. The highest important elevated coral 
reef rocks occur at an altitude of about 35 feet above sea level off the 
sides of the stream mouth. The stream has cut and maintained a gorge 
through about 165 feet of Umestone and marl which are topographi- 
cally above the reef and which are not coral reef rocks, but which 
are bedded and were formed by other agfencies. Other instances of 
these relations might be given. 

The conditions around the Habana Harbor are interesting in this 
connection. Limestone of upper Oligocene or Miocene age occurs 
at the Morro and forms the h^her land along the shore east of the 
city, and it outcrops at lower altitudes in the western part of the 
city; but the drainage at the south end of the harbor has cut through 
the limestone and exposed the underlying rocks, serpentine, rotten 
diorite, etc.; and that underground solution is active is indicated by 
the presence of springs along the serpentine contact. The condi- 
tions are here favorable for erosion by both mechanical cutting and 
solution in the area lying behind, while a channel has been main- 
tained across the limestone on the sea front. This basin after it 
was outlined was submei^ed. 

It is intend^ to give a much fuller discussion of the Cuban harbors 
in a paper now almost ready for press. The differences in form, and 
ihe causes to which the differences are due, are worthy of far more 
detailed treatment than is practicable in this place. I will end this 
part of the present discussion by saying that corals have in certain 
instances played a subordinate r6le by narrowing the mouth of a 
harbor and by preserving a constricted outlet. That the outlets of 
the basins here considered were constricted by reef rocks, now ele- 
vated, is shown by the conditions in Habana and Santiago harbors, 
and that similar constriction is now taking place by similar agencies 



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286 BULLETIN 103, UNITED STATES NATIONAL MUffiUM. 

is exemplified in many of the West Indian Islands. As the coral 
rock is usually harder than the rocks on which it rests, after its 
emergence it protects the narrow exit behind which erosion is more 
rapid and enlarges the basin. 

From the remarks already made it appears unnecessary to discuss 
specially which are the older — the drainage basins occupied by the 
harbors or the coral reefs now elevated about 30 feet. However, 
that the Santiago basin is older than the coastal soborruco is shown 
by finding the soborruco within the harbor mouth; and as I found 
recent species of reef corals, apparently in place, on the east side of 
Habana Harbor, south of the Monro, at a height of 30 feet aboYe sea 
level, the 30-foot reef seems to extend into the mouth of Habana 
Harbor. * The valleys are clearly older. On page 264 of this paper a 
special point was made of the unconformity between the elevated 
Pleistocene reefs and the underlying Miocene material and the in- 
ference was drawn that the reefs were formed during subsidence after 
erosion of the basement under them. This is precisely the interpreta- 
tion Professor Davis had made of the relations in the elevated reefs 
of the New Hebrides, but it seems such relations may develop in the 
same cycle, and, in my opinion, they are of slight importance in their 
bearing on the general theory of coral-reef fc^rmation. 

The Isle of Pines furnishes important information on changes in sea 
level aroxmd Cuba. This island is nearly opposite Habana, 60 miles 
south of the south coast of Cuba, from which it is separated by water 
less than 10 fathoms deep. It comprises two parts, a southern which 
is mostly swamp, and a northern which is topographicaUy higher. 
The surface of the northern division is mostly a plain, really a 
peneplain (see pi. 72, fig. A), above whose surface stand monadnoeks 
of harder rocks (pi. 72, fig. B). This island is very different from 
the main island for, as no Tertiary or Cretaceous marine deposits 
are known to occur on it, it appears to have r^siained above sea 
level during these periods, but it has experienced the later changes 
of sea level which affected the larger island and during Pleistocene 
time it was joined to Cuba. The peneplain was formed at a lower 
level than that at which it now stands, it was then sufilciently uplifted 
to permit streams to cut into it, and has then been depressed, 
thereby drowning the mouths of the streams, but not bringing the 
plain surface so low as it formerly stood (pi. 72, fig. C). The 
coast line of the Isle of Pines and that of Cuba immediately north of 
it both are indented by the embayment of stream mouths throu^ 
geologically recent submergence. 

That the Isle of Pines was joined to Cuba duriag Pleistocene time 
is shown convincingly by its land fauna. Every species of reptile, 
except one, found on it. Dr. L. Stejneger informs me, is known to 
occur in Cuba, and two species of the mammaUan genus Capromys 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 287 

are common to both. Dr. Paul Bartsch tells me that the Isle of 
Pines is only "a chmik of Cuba" and that its land Mollusca represent 
a faunal area as closely related to the faimal areas of Cuba as are 
the different faimal areas in Cuba to one another; that is, faimaUy, 
the Isle of Pines is simply a portion (a faunal area) of Cuba. There- 
fore, it is clear that the Isle of Pines has been severed from Cuba in 
the latest Pleistocene or Recent geologic time. 

Practically all the Cuban shore line has now been considered except 
that on the north side of the Province of Pinar del Rio, within the 
Colorados Reefs. Guadiana Bay is a nearly typical estuarine em- 
bayment, while sUghter embayment of other stream mouths is 
indicated, and lines of islands extend seaward from some headlands. 
The shore line clearly indicates submergence. Mr. J. B. Henderson 
and Doctor Bartsch, however, tell me that there is positive evidence 
of minor uplift west of Guadiana Bay. * 

The Cuban shore line as a whole shows evidence of Recent oi; latest 
Pleistocene submergence, and this submergence has influenced the 
modem coral-reef development. 

Regarding the amoimt of Recent submergence of the Cuban shore 
line, reference to the table on page 282 shows that' there is close ac- 
cordance in the depths of the channels or harbors, except certain 
ones that will be discussed later. These indicate that prior to the 
last submergence the land stood about 100 feet or slightly more, 
about 20 fathoms, higher than at present. The amoimt of emergence 
would establish a broad land connection with the Isle of Pines. 

The discrepant harbors are Nuevitas Bay, which shows an excess 
of only about 27 feet, Nipe and Tanamo bays, and the channel 
leading from Livisa and Cabonico bays, on the north coast, and 
Cienfuegos on the south coast. The harbors with the discrepant 
depths on the north coast all occur on the north side of the Province 
of Oriente and at the eastern end of the Province of Camaguey. 
They seem to indicate deeper submergence than at other places and 
that the submergence has not been uniform in amount for the entire 
coast. However, the depths do not contradict a Recent rise of sea 
level to an amount of about 20 fathoms. The harbor of Cienfuegos 
would be expected to be abnormal, for the fault line which runs 
northward from Cape Cruz intersects the shore line at its mouth 
(see text-fig. 15). It is possible that structiual relations have also 
influenced the depths in the other harbors and channels that are 
discrepant. Regarding these it will be said that except Nuevitas 
Harbor they occur within a linear distance of 31 miles. Nipe Harbor, 
the westernmost of the group, lies on the north side of Loma de 
Mulas, while it, Livisa, Cabonico, and Tanamo harbors all are on 
the north side of Sierra Cristal. 

> Henderson. J. B., Cruise of the Tomas Bantm, pp. 161-164, New York, 1916. 



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288 BULLETIl^ 103, UNITED STATES NATIONAL MUSEUM. 

The great extent and relatively uniform height of a coral-reef 
terrace between 30 and 40 feet above sea level favors the interpreta- 
tion that the geologically Recent shift in position of strand line has 
been without pronounced crustal deformation. 

The relations of the oflF-shore reefs to the platforms on which they 
grow will now be briefly considered. A detailed description of the 
reefs is unnecessary here, as it would be only a repetition of that 
already given by A. Agassiz ^ and the accounts contained in the 
West Indies Pilot.* It need only be stated that the best developed 
off-shore reefs on the north coast are the Colorados Reefs, between 
Bahia Honda.and Cape San Antonio; and that off the south coast 
the best are those between Trinidad and Cape Cruz and those east 
and west of the Isle of Pines. Mr. John B. Henderson has devoted 
attention to the Colorados Reefs in his ''Cruise of the Tomiu 
Barrera.^^ Have the reefs off the south coast grown up on the 
surface of preexistent platforms or are the platforms due to infilling 
behind a reef during subsidence ? 

The area between Trinidad and Cape Cruz wiU be considered 
first. The fact that the reefs form disconnected hillocks or mounds, 
sometimes of mushroom shape, above a plain surface, which in places 
is 50 miles wide along a line perpendicular to the shore, while on 
the seaward side of the reefs there are large areas of shallow platforms, 
without any margining reefs, seems conclusive evidence against the 
platform having been caused by infilling behind reefs. 

The following, in my opinion, is the correct explanation: The Ut- 
toral geologic formations from Cape Cruz to Trinidad are mostly 
upper Oligocene or Miocene marls and limestones which dip under 
the sea at relatively low angles. They dip into the Cauto Valley, 
which is a gently pitching syncline, and into its seaward continua- 
tion, the Gulf of Guacanayabo. The embayment northeast of Boca 
Grande passage is probably abo synclinal in structure. The abrupt 
undersea termination of the platform is most reasonably explained 
by a submarine fault which runs from Pimta SabaniUa, at the mouth 
of Cienfuegos Harbor, to Cape Cruz. The coral reefs have grown 
up on the surface of a plain underlain by geologic formations that 
were gently tilted seaward and faulted along the line indicated. 

That the Isle of Pines was joined to Cuba during Pleistocene time 
has, I beUeve^ been shown in a convincing manner. As the Miocene 
and upper Oligocene formations from Batabano to Pinar del Rio dip 
under the sea at low angles they must underUe the flat bottom of 
the Gulf of Batabano. That the submarine slope from East Guano 
Key to off Cape San Antonio is determined either by a fault or by a 
very steep flexure is clearly indicated, as off the south shore of the 

1 Bull. Mus. Comp. Zool., vol. 26, pp. 133-136, 1804. 

s West Indies PUot, vol. 1, pp. 199-332, 1913 (U. S. HydrograpUc Office). 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 289 




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290 



BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 



Isle of Pines the descent from the shore in 9 sea-miles is 13,080 feet, 
a slope of about 1 in 4. The Gulf of Cazones appears to have be^i 
outlined by faulting. This shelf differs from the one considered in 
the preceding paragraph, in that the Isle of Pines, whose area is 
about 1,^00 square miles, stands on its outer mai^in, and apparently 
has affected the course of the fault. However, there was here also 
an undersea flat, which was produced by the gentle seaward tilt of 
low-lying geologic formations, and its outer margin was also deter- 



lS8ft 







0C. ConiMttt 



L.L.POftTCS CO^ Kfc 



FZQ. 16.— Chabt of oolobados KXKrs, CUBA. From U. 8. htdsooraphic chart no. 966. 

mined by faulting. The living reefs are growing on its submerged 
unfaulted part, above which they rise as disconnected patches or as 
a broken barrier. 

The Colorados reefs (text-fig. 16) grow as patches of barrier reefs 
or upon a shelf, which, according to Henderson, largely consists of 
coral rock that had been uplifted above the sea and then depressed.^ 

The conditions imder which the Cuban offshore reefs are growing 
can be very easily summarized, as follows: (1) They are superposed 

1 Henderson, John B., Cruise of the J\ma3 Barren, pp. 62-64, 126-190, 1916. 



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QEOLOOY AND PALEONTOLOGY OP THE CANAL ZONE. 291 

on flats submei^ed in geologically Recent time; (2) the amount of 
the submergence of Cuba was about 100 feet. 

BAHAMAS. 

Alexander Agassiz has in his reconnaisance of the Bahamas^ the 
following very significant statement: 

May we not to a great extent measure the amount of subsidence which must have 
taken place at certain points of the Bahamas by the depth attained in some of the 
so-called ocean holes, as marked on the charts? Of course we aasiune that they were 
due in the aeolian strata to the same process which has on the shores of many islands 
formed potholes, boiling holes, banana holes, sea holes, caverns, caves, sinks, cavities, 
blowholes, and other openings in the aeolian rocks. They are all due more or less to 
the action of rain percolating through the aeolian rocks and becoming chaiged with 
carbonic acid, or rendered acid by the fermentation of decomposed vegetable or ani- 
mal matter or by the action upon the limestone of sea water or spray under the most 
varying conditions of elevation and of exposure. None of them have their upper 
openings below low-water mark, though some of them may reach many feet below 
low- water level. Ocean holes were formed in a similar way at a time when that part 
of the bank where they exist was above high-water mark, and at a sufficient height 
above that point to include its deepest part. The subsidence of the bank has carried 
the level of the mouth and of the bottom of the hole below high-water mark. 

From the description of the strata which crop out upon the banks in the vicinity of 
some of the ocean holes at Blue Hole Point, there seems to be little doubt that the 
stratification characteristic of the aeolian rocks has been observed. 

The deepest of the holes mentioned by Agassiz has a depth of 38 
fathoms, *'in the extension of the line of Blossom Channel leading 
from the Tongue of the Ocean up on the bank." 

I have had opportunities to study such *' holes" or solution wells, 
above sea level in Florida and have examined many of them, both 
above and below sea level, in the Bahamas. Mr. E. W. Forsyth 
sounded other * ' holes " and reported the results to me.' The depths of 
the holes range from about 2 fathoms tq as much as 33 fathoms, the 
deepest hole in Fat Turtle Sound, North Bight, Andros Island, 
sounded by Mr. Forsyth. As in my opinion Agassiz's deduction as 
to the origin of these holes is incontrovertible, they indicate a stand 
of the land during Pleistocene time at least 228 feet higher than at 
present. Shattuck' and Miller accept a higher stand of 300 feet, 
followed by submergence of 300 feet, and conclude that this move- 
ment in strand-line position was followed by emergence, to an amount 
between 15 and 20 feet. From my own experience in the Bahamas 
the last change in the position of strand line was accompanied by 
minor differential crustal movement. For instance, at Nicollstovn 
Light, Andros Island, a sea cave stands at such a height above the 
sea as to show conclusively an elevation of 18 feet above the position 

1 Mas. Comp. Zool. BuU., vol. 36, pp. 41-43, 1»4. 

s Vaughan, T. W., CanMgto Inst. Wasfalnfton Tearbook No. 18, p. 230, 1916. 

• Shattuck,.0. B., and lflJI«»r, B. L., Physiography and geology of tho Bahama Islands, The Bahama 
Islands, pp. 10, 3D, 1006w 



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292 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

at which it was formed; but 4,000 feet southeast of the cave the 
elevation is only about 4 feet in amount. I have given more infor- 
mation on this minor uplift in the paper referred to in the footnote.* 

Agassiz, Sbattuck, and Miller, and I agree as to the geolc^- 
oally Recent submergence of the Bahamas. 

Tte accompanying diagram (text-fig. 17) indicates the relations 
of the barTiei" reef off the west side of Andros Island to the platform 
on the edge of which it is growing. This reef is growing on the edge 
of a platform that had stood above sea level at least as much as 192 
feet. It was perforated by solution wells and then submei^ed. . The 
perforations in the platform show that it antedates the barrier reef, 
and that its formation is not dependent on agencies associated with 
the presence of the reef. There is here another instance of a reef 
formed during or after submergence, and superposed on the surface 
of an antecedent platform. 

Sk)' oolite 
^ we n^dgg Coral reef SMlifiXtf. 

Libmerged Terrace i^fWift. 

■ ooivnion wens o-z^/xiaunirno ^ 

I deep through oolite 

2001J I 

Solution well 31-33 fathomd deep 



4001 



eo<d 



Horizontal acala 
o tjyo Afiop 




L-JIOOfintt. 



FlO. 17.~DU0RAMMATIC 8£CTI0N ▲CE088 THS BABBIBK BSIf, ANDBOS ISLAND, BaHAILLS. 

The relative importance of the constructional rdle of the living 
reef will be briefly mentioned. The Pleistocene oolite of the Bahamas 
is not coral-reef rock, as was contended by A. Agassiz. It is composed 
of calcium carbonate chemically precipitated on extensive submarine 
flats.^ I have several times published the estimate ^Hhat on Andros 
Island, Bahamas, the ratio of the constructive work of the present 
reef to that of agencies that previously resulted in the formation of 
the Pleistocene ooUte is approximately as 1 to several thousand, or, 
as a constructive agent, chemical precipitation has been several 
thjousand times more effective in forming hmestone than corals."' 

Before passing to the discussion of the next area it should be 
pointed out that the amount of submergence of the Bahamas, 228 

1 Carnegie Inst. Washington Yearbook No. 13, p. 229, .1915. 

s For the most recent dlsonssions of this sul^eot, see Vauglian, T. W., Some shoal-water bottom samples 
from Munay Uaad, etc, Camegie Inst. Washington Pnb. 213, pp. 277-280, 1918; Qiemloal and organic 
deposits of the sea, Geol. Soc. Amer. Bull., vol. 28, pp. 983-944, 1918. 

« Wash. Acad. Bd. Joum., vol. 4, pp. 26, 27, 1914; Caniegie Inst. Washington Pnb. 213, p. 279, 1917. 



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GEOLOGY ANI> PALEONTOLOGY OF THE CANAL ZONE. 293 

feet, is greater than that, about 120 feet, indicated for the areas 
abeady considered, unless the notches in the outer edges of the SU 
Martin Plateau and the Virgin Bank really indicate a position of sea 
level 40 fathoms lower than present sea level. 

BERMUDAS. 

Alexander Agassiz has given a good aocount of proto-Bermuda, that 
is of the extent and general physical character of the Bermudas pre- 
vious to the submergence that has left the group in very nearly the 
form in which we now know it.*- Recently Prof, L. V. Pir^son has con- 
tributed two highly valuable articles to the literature on the geology 
of the islands, basing his interpretations largely upon a study of 
samples from a well bored in Southampton Parish, on the slope of a 
hill about a mile west of the lighthouse on Gibbs Hill, from a height 
of 135 feet above sea level to a depth of 1,413 feet below the surface, 
or to a depth of 1,278 feet below sea level.^ 

There were penetrated in the weU mentioned three major classes 
of material, as follows: (1) From the surface to a depth 383 feet 
below it, limestone; (2) from 383 feet to 600 feet, oxidized volcanic 
material; (3) below 600 feet to 1,413 feet, with one slight exception, 
basaltic, usually black lava. Pirsson concludes the first of his two 
articles with the following statement: 

It appears to the writer that what has been learned regarding the history of the 
Bennuda volcano has an important bearing on the question of the way in which the 
platforms on which coral islands, barrier reefs and atolls are situated, have been 
fOTmed. There is of course nothing new in the idea that these may be volcanic in 
origin, only in Bermuda we have once more a positive demonstration of the foct. We 
have also seen that, provided the volcanic masses are of sufficient antiquity, they 
may, even though of great size, have been reduced to sea level, furnishing platforms 
of wide extent. As mentioned above, such masses reduced to sea level would con- 
tinue to project from the ocean abysses indefinitely and many of them may be of great 
geologic age. There is nothing in the mere size of any of the atolls of the Pacific which 
would preclude their being placed on the stumps of former volcanic masses; it is not 
intended to assert by this that the foundation in every case is necessarily a volcanic 
otie. If such masses have once been brought down to sea level and continue to exist 
and that level changes within limits from time to time by warpings in different places 
of the sea floor, or by an accumulation of ice on the lands and its melting, as suggested 
by Daly, then conditions of shallow water over them may be established suitable for 
their colonization by those organisms concerned in the production of the so-called 
coral reefe, which may be formed under the conditions postulated by Vaughan. 

It was the imderstanding between Professor Pirsson and me that 
I should prepare a report on the calcium-carbonate samples. The 
following is a preliminary statement, accompanied by determinations 
of the Foraminifera by Dr. Joseph A. Cushman. 

> Agaasis, Alexander, A visit to the Bermadas in Iforoh, 1894, ICns. Comp. Zool. Bull., vol. 36, pp. 273- 
3n, pi. 3, 1806. 

> Pinson, L. V., Geology of Bennnda Island, I. The igneous platform, Amer. Joom. Sd., ser. 4, vol. 38, 
pp. 180-aoO; n. Petrology of the lavas. Idem., pp. 331-344, 1914. 



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294 



BULLETIN 103, UKITBD STATES NATIONAL MUSEUM. 



Preliminary deicription of the limutone tamplee and litt qf spedee of Foramin\ferafnm 

the Bermuaa weU, 



No. of speoimen 

andaepth 
below surface. 



Desoriptioii. 



Species of FonmlnifBra. 



1 ((MIfeet) 



.2 (61-110 feet) 



8 (110-216 feet) 



4 (216-241 feet) 



6 (241-286 feet) 



6 (286^1 iB6t) 



7 (331-841 iset) 



8 (841-383 feet) 



Ught croam-oolored limestoiie; mixture of oal- 
ate and aragonlte: most of the oonstitiieBt 
particles angular; largely or mostly broken 
remains of organisms; occasional small round 
grains 0.10 mm. or lees in diameter, may be 
agnegates of chemically predpitated calcium 
carbonate. Largely or mostly an organic 
limestone. 



Light cream-colored limestone; mixture of cal- 
dteandi 



0.04 mm. or less in diameter, may be aggre- 
gates of chemically precipitated material. 
Iiargely or mostly an organic limestone. 

Li|dit cream-colore<{ limestone: mixture of cal- 
ate and aragonite, apparently but little ara- 
gonite; laigelv a reorystalllsed limestone, 
without conspicuous grams; some small pock- 
ets contain pulverulent calcium carbonate; 
some pieces eranular. A few grains 0.05 to 
0.8 mm. in diameter resemble small oolite 
irrains. The rock is mostly a foraminiferal 
limestone, the Foraminil»ra embedded in a 
cryptocrystalline matrix. 

Whitish limestone, Very slight yellowish tinse, 
some blackish particles; mixture of acagomte 
and calcite ; speoimen consists mostly of broken 
rock fragments; an occasional small pebble, 
one 2.5 nmi. as maximnm diameter; constit- 
uent material largely oreanio,ForaminilBra. 
fragments of moUusks, shells, etc. Most small 
particles angular: a few less than 0.12 
nmi. appear oolitic. One 0.00 by 0.17 mm. 
in site had form of an oolitic ellipsoid. Mostly 
an organic limestone. 

Whitish, faintly yeUowlsh, nulywulent lime- 
stone: mixture ofcalcite and aragonite. 0>m- 
paratiyely few tests of organisms, some Fora- 
minifera, many small rounded nrains and 
cryptocrystalline material. Some of the round 
grams appear oolitic; one of these is 0.11 by 
0.15 nun. in sixe. It appears that a consider- 
able proportion of this bed Is a chemical pre- 
cipitate. 

White, pulverulent limestone; mixture of cal- 
cite and aragonite. No organic tests were ob- 
served. Round grains up to 0.1 or 0.2 mm. 
appear to be ooUte; small round grains 0.04 
mm. in diameter. Much cryptocrystalline 
material. This bed appears to be largely a 
chemical precipitate. 

White, friable limestone; mixture of calcite and 
aragonite. Round grains which range in di- 
ameter from 0.22 to 0.45 mm., may be oolitic. 
Small grains, 0.09 mm. in diameter seem defi- 
nitely oolitic. Besides the rounded, there are 
broken angular gndns and much cryptocrys- 
talline materiaL Few or no orgsnio tests. 
This appears to be largely a oheimcal precip- 
itate. 



Lightoolored, earthy, 



impure 
[y calcite. 



limestone; some Iron pyrites; mi 
if aragonite is present the proportion is smallc 
ManyForaminifera, Jfufiini«me«, fragments of 
coral. Bryosoa, etc.: many rounded grains 
which may be detntal; no definitely oolitic 
grains were observed. A thin section shows 
many Foraminifera embedded in a cryptoorys- 
talUne matrix. This bed is an Impure, forami- 
niferal, shoal water limestone. It may con- 
tain some chemically precipitated material. 



TextHJaria agglvUmnB d'Otb 
Polifatomdia 9triatopunemm 

AMoll. 
Polyttiomdla species. 
A mpMHtgin* U9$onH d'OrUgny. 
QuinfuOoctUinm retkutata d'Orbigiiy. 
O. obtonoa Montasu. 
Q. oubenafM d'Oroimy. 
PeneroplU penutu* FbrskU. 
OrNctt/iiia otftMcs Fiohtel & MoO. 

Textutaria OfgluHnmns d'OrUmy. 
PolifStomeUa <lriatop»ncteCB ^olital 

& Moll. 
Polfttomau spedM. 
A mplUHegina U»wiUi d'Orl 
CtuinmOoeuUna reticulata & 
6. auberiana d'Orbigny. 
OrbUMlina adunea n^tal A M61L 

CUvuUfM anfutarU d'Orbigny. 
Planorbulinm larvata Parker& Jones. 
TVunoetuZtna spedee. 

'^^sr ""'^^^ »*« 

Ampkiiteifina letionii d'Orbigny. 
Truoeulina cL T. draitemBome- 



OfiOeuUna adrnnet Fiohtel A M6IL 



TnMoUulhM spwAes, 
PuMimUwt eanarientiM d'Orbloif . 
Polpttomeila ttrtatopunelata FicAti 

&MoU. 
PotyttomeOa species. 
A mpkittegifia UnonH d'Orbigny. 
TTiioculina linneana d'Orbiny. 
Orbiaainmudunca Fiohtel A MolL 



BoUslna suedes. 

TVtmoaAtaiM species. 

DiteorbU pUariiboana d'Orbigny. 

.i mpMsM^na species. 

QtUnquOocuUna reticuiata d'Orbigny. 

BUoc '• 



None reported. 



Amphitteffina species. 



JVttinfiittUtet species. 



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GEOLOOT AND PALEONTOLOGY OP THE OANAL ZONE. 295 

This examination reveals three kinds of Umestone, the uppermost 
of ivhich subsequently may be subdivided. The three divisions are 
as follows: 

Specimens 1-4 (0-241 feet) represent a limestone which is largely or 
mostly of organic origin, but which may contain a few grains of 
chLemically precipitated material. This corresponds to the upper 
f aiinal division recognized by Cushman. 

Specimens 5-7 (241-341 feet) represent a pulverulent limestone, 
composed of rounded grains imbedded in finely crystalline material. 
The grains in their size and shape resemble oohte, and some grains 
showed with greater or less distinctness suggestions of oohtic struc- 
ture. The foraminiferal fauna is meager, but it differs from that of 
specimens 1-4 and the imderlying bed represented by specimen 8. 
It seems safe to draw the inference that this division of the lime- 
stone is in part, ^t least, a chemical precipitate. 

Specimen 8 (341-383 feet) represents an impure, foraminiferal, 
earthy limestone, or a calcareous marl, in wbich there may be some 
chemically precipitated material. This bed is the uppermost in 
which the Nummvlites reported by Cushman occur. It was also 
fotind in the underlying bed No. 9, 383-393 feet. 

Probable geologic age of the limeetone in the Bermuda well, 

[Height of well moath above sea level, 135 fset.] 
Samplee. Probable geologio age. 

Ytixtn. 1-241 feet Recent and Pleistocene. 

From 241-286 feet Pliocene or Miocene. 

From 331-341 feet Nothing determinable. 

From 341-393 feet Oligocene or £k>cene {Nummuliies), 

FroxQ 393-485 feet Eocene? (no Nummvlites), 

At> outline of the geologic history of the Bermudas subsequent 
to the volcanic activity seems to be as follows: 

I>octor Cushman's identification of the Foramimfera from the 
Bermuda well shows the presence of an imdetermined. species of 
PolystomeUa between 393 and 480 and between 480 and 485 feet. 
These depths are well down in the oxidized zone and indicate marine 
conditions which persisted throughout the deposition of the super- 
incumbent material. Other Foraminif era occur between 383 and 393, 
one of them being a species of NummuLiteSj which was also obtained 
from the basal bed of limestone at a depth of 341 feet. As the 
genus Numw/alites is, according to our present knowledge, confined 
to the upper Ek)cene and Oligocene formations in the southeastern 
United States and the West Indies, the inference may be drawn that 
the Bermuda samples between 341 and 393 feet probably represent a 
geologic formation of either Eocene or Oligocene age, and that those 
from 393-486 feet represent a formation of probably Eocene age. 
87149— l»—BuU. 103 8 



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296 BULLETIN lOSy UNITED STATES NATIONAL MUSEUM. 

Until the specimens of Nummulites from the Bermuda well hare 
been identified with species of known stratigraphic position a more 
definite statement can not be made. It appears safe to assign an 
Eocene or pre-Eocene age to the Bermudian volcanic activity. 

The calcareous sediments, therefore, b^an to accumulate on a 
submerged volcanic basement in ESocene or lower Oligocene time, 
and the submergence progressed until the basement, in probably 
Miocene time, was entirely blanketed by calcareous deposits 100 
feet thick, which differ in their physical aspect both from the under- 
lying nummulitic rock and the overlying organic limestone. This 
rock is probably in considerable part a chemical precipitate. The 
well samples indicate no stratigraphic break at either its top or it 
base. 

The limestone from a depth of 241 feet to the surface is a shoal- 
water, organic deposit, in which living species of Foraminifera are 
abundant. Its age is probably Pleistocene, although the lower part 
may prove to be Pliocene. The shoal-water nature of the limestone 
indicates continued slow subsidence. 

The subsidence which apparently had been interrupted by no 
period of emergence since Oligocene time was succeeded in Pleisto- 
cene time by uplift to an amoimt of probably more than 100 feet. 
AU the surface rock of the Bermudas except some in areas of low 
elevation is considered by the geologists who have visited the islands 
to be eolian deposits. However, certain of the published illustra- 
tions suggest that in some exposures there are in the bedding hori- 
zontal planes intersecting the inclined layers. Oross-bedding between 
horizontal planes is a structure characteristic of shoal-water or beach 
deposits but not of eolian deposits. A more critical study of the 
bedding of the Bermudian rocks may discriminate elevated cross- 
bedded water-laid and eolian deposits. However this may be, ihe 
period of uplift under consideration was the time of the Greater Ber- 
muda, which has been admirably described by WilUam North Rice, 
A. Agassiz, and A. E. Verrill. According to the latter, the area of 
Greater Bermuda was somewhat more than 230 square miles, or about 
11 times that of the present land surface, which is estimated as hav- 
ing an area of 19i square miles.^ The evidence indicates that the 
elliptical area inclosed by the outer reefs wasentirely above sea level, 
as perhaps also were the surfaces of Challenger and Argus banks. 

The last important change in the relations of sea level was, as 
Verrill has so ably shown, submergence to an amount of about 100 
feet, reducing the land area from that of 230 square miles during the 
period of Greater Bermuda to that of 19^ square mUes, the present 

i Conn. Acad. Arts and Sd. Trans., vol. 12, p. 52, 1905. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 297 

area. The evidence is not decisive as to there having been a slight 
emeigence, of 6 to 10 feet, since the great submergence. 

As Verrill has shown, the Bermuda limestone is composed not of 
coral d6bris, except in a subordinate proportion, but is made up of 
broken, more or less triturated, calcareous tests, largely of moUusks. 
He designates the material as ''shell sands.'' The Bermudas are, 
therefore, inappropriately called "coral islands." The recent corals 
are growing on a foundation of older lime rock, brought into its 
present relation to sea level by sulvnergence. 

In that the last dominant change in the position of its strand line 
was by submergence, Bermuda accords with the Florida coast, the 
Bahamas, Cuba, and most of the smaller West Indian islands. 

FLORIDA. 

Strand-line oscillation in Florida has attracted the attention of 
many geologists, among whom may be mentioned Shaler, Heilprin, 
and Dall of the earlier investigators, and Matson, Sanford, Sellards, 
Shaw, and myseK of the later ones, ^haw and I have recently 
reviewed the subject.* That subsequent to formation of the Pleis- 
tocene barrier reef of Florida, the reef tract was elevated to a height 
about 50 feet above its previous stand and that this elevation was 
followed by submergence to an amount of about 30 feet is shown by 
(1) a submerged cave at Miami; (2) subnierged solution well below 
sea level, near East Bahia Honda Key; ' (3) submerged peat bed at 
Key West; (4) submerged indurated, cemented, recrystiJlized oolite 
under the Marquesas; (5) submeiged wave-cut terrace front at 
Tortugas. 

In addition to this evidence Shaw and I say in the paper cited: 

Additional deductioDB of importance may be made from the submarine physiography 
at depths beyond 10 fathoms. Although the investigations are at present only in a 
preliminary stage, it may be said that along the sides of the Gulf Stream from opposite 
Miami to Satan and Vestal Shoals, just west of Sand Key, the Coast and Geodetic Survey 
charts indicate Mrly uniform slopes from 10 to 100 fathoms, but there may be narrow 
terraces which are not brought out by the soundings. West of Vestal Shoal the sea 
bottom drops suddenly from 10 to 20 fathoms, with a flat or gently sloping surface 
between 21 and 28 fathoms. South of Goalbin Rock there is an escarpment between 
10 and 30 &tthoms, a flat or gentle slope between 30 and 40 fathoms, and another flat 
or gently sldping area between 40 and 50 fathoms. The soimdings are not sufficiently 
niunerous to trace surfaces with a feeling of confidence, but the scarp from 10 to 
between 25 and 30 fathoms is clear cut and can be followed for 25 miles to the west end 
of the Quicksands. Westward in the vicinity of Tortugas there are, besides, the 
bottom of Tortugas lagoon and the surface of the shoal 7 to 10 miles west of Loggerhead 
Key, two undersea terrace plains, one at a depth of 15 to 17 &tthoms, the other, which 
is a large plain west of Tortugas; ranges in depth from 28 fathoms on its landward to 
36 tAihoJDB on its seaward edge, and has an east and west width of 10 miles. The 15 
to 18 tiithom flat is especially well developed south and southwest of Tortugas. It is 

1 Vsoghan, T. W., and Shaw, E. W., Qeologlo Investigations of the Florida coral-reef tract, Carnegie 
Inst. Washington Yearbook No. 14, pp. 332-388, 1010. 
t Oral oommnnicatSon of Mr. Sanuxel Sanford. 



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298 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

separated by a scarp from the 28 to 36 fathom flat, and by another scarp from the 
shallower levels in Tortugas. The presence of the continuous scarp from Goalbin Rock 
to off the west end of the Quicksands, with a depth of 25 to 30 fathoms at its foot, and 
the presence of a terrace 28 to 36 fathoms deep, 10 miles wide, and bounded on its land- 




Fio. 18.— Chabt or nobthkbn end or rLOBunAN barrebr RXKr. From Unrbd SfATss coast amd 

GEODETIC SURVEY CHART MO. 166. 

ward margin by a similar scarp, suggest that the portion of the Florida reef tract west 
of Key West at one time stood some 20 fathoms higher than now, while the 15 to 18 
fathom terraces suggest another, shallower stand of sea level. 

Although the tracing of the oscillations of the Florida reef tract can not now be 
made in detail, it seems probable that it at one time stood more than 120 feet higha 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 299 

than at present (land has been submeiged to a similar amount). Besides the suggested 
larger swing there have been intermediate stands of sea level and niunerous minor 
oscillations. The last movement of importance was one of submergence, but subset 
quent to it there has been a minor uplift of some 10 feet or slightly more in the vicinity 
of Miami. * 

The accompanying figure (fig. 18) shows that the flat that the living 
barrier-reef margins or aboye which coral-reef patches rise extends 
beyond the northern reef limits, near Fowey Rocks. The living bar- 
rier reef has developed seaward of the Pleistocene barrier near the edge 
of a previously prepared platfwm, for the continuity of the platform 
irrespective of the presence of the reefs shows that its origin is inde- 
pendent of them. 

CAMPECHE BANK. 

The best known reef on the Campeche Bank is Alacran Reef, which 
was described by A. Agassiz in considerable detail in 1888.* (See 
pi. 73, photograph of model.) Heilprin in 1891 ^ said regarding 
Yucatan, "the evidence is all but conclusive that there has been 
recent subsidence"; but I am unable to discover in his article the 
basis of this opinion. Dr. C. W. Hayes oraUy informed me shortly 
before his deeply lamented death that there is clear evidence of 
recent submei^ence aroimd Terminos Lake at the base of the penin- 
sula on its west side. The lagoons between Progreso and Holbox 
Island are strongly suggestive of submergence. There is a steeper 
slope between about 20 and 28 on the outer edge of the bank, indi- 
cating change in position of sea level by submergence, similar to the 
change already recorded for St. Thomas and other West Indian 
islands. 

In this connection the following quotation from Alexander Agassiz 
will be introduced :* 

In fact, what I have seen so to in my exploration of the coral reefs of the^West Indies 
would show that wherever coral reefe occur, and of whatever shape, they form only a 
compaiativdy thin growth upon the underlying base, and are not of great thickness. 
In Florida they rest upon the limestones which form the basis of the great peninsula. 
On the Yucatan Bank they are underlain by a marine limestone. In Cuba they abut 
upon the Tertiary limestones of its shore. Along Honduras, the Mosquito Coast, and 
the north shore of South America they grow upon extensive banks or shoals, parts of 
the shore plateau of the adjoining continent, where they find the proper depth. 

I doubt if there is any one bold enough to claim that Campeche 
Bank has been formed by infilling behind a barrier reef, for it is too 
obviously due to a large gentle flexure of the earth crust or some 
other kind of broad structural uplift, and that in suitable places 
coral grows on the surface of the submarine plateau formed in the 
manner indicated. E. W. Shaw* collected a few bottom samples 6 to 

I Agassiz, A., Three crnlses of the Blake, vol. 1, p. 71, 1888. 

* HeUprtn, A., Qeologtoal researches ih Yucatan, Phila. Acad. Nat. Sd. Proc. for 1891, p. 148. 
i Mus. Comp. Zool. BulL, vol. 36, p. 173, 18M. 

* Shaw, E. W., Oral oommanicatlon. 



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300 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

8 miles off shore at Progreso^ and in these he found only two fragments 
of coral, the main mass of the samples being shell fiagments. 

HONDUBAN REEFS. 

Although this is an important barrier reef, its length 1[>eing 125 
sea-miles, I know of no adequate published description ot it, nor of 
any published accoimt of the shore line or of the oscillations of the 
strand line behind it. The configuration of Honduras Bay and of 
the Gulf of Dulce, which lies inland from it and is connected with it 
by a waterway, as well as that of Chetumal Bay, points dearly to 
submergence. The reef occupies the outer edge of a platform 10 to 
22 miles wide ^nd is separated from the shore by a channel from 11 
to 33 fathoms deep. This is a remarkably continuous barrier reef, 
but it shows discontinuity at its southern end and therefore evidence 
of superposition. 

MOSQUITO BANK. 

Hayes, although he was not giving particular attention to coral 
reefs, has made one of the finest studies of a shore line in a coral- 
reef area as yet published.* The following is quoted from his 
article:' 

7. In middle Tertiary time the region was elevated and subjected to long-continued 
flubaerial degradation, and the narrower portion of the iBthmus was reduced to a pene- 
plain, with monadnocks at the divide near the axis. There is no evidence that open 
communication has existed between the two oceans across this portion of the isthmus 
since the middle Tertiary uplift. 

8. In post-Tertiary time the region was again elevated and the previously developed 
peneplain deeply trenched. 

9. A recent slight subsidence has drowned the lower courses of the river valleys, 
and the estuaries thus formed have subsequently been filled with alluvial deposits. 

J. E. Spurr furnished me a note* confirming Hayes's deduction 
regarding the submei^nce of the lower courses of the streams on 
the east coast of Nicaragua. Subsequently I had profiles drawn 
across Mosquito Bank (see text fig. 11, page 275).^ These indicate 
submergence to an amoimt of about 20 fathoms. As on Mosquito 
Bank there is a submerged terrace front between about 20 and 25 
fathoms in depth, the bank had to exist previous to formation of 
that feature, and as the living reefs grow on the shallower flats, 
which according to available evidence was out of water during at 
least a part of Pleistocene time, they are necessarily superposed on 
an antecedent basement. Furthermore, the enormous area of the 
flat and the relatively small areas occupied by living reefs, lead to 
the same conclusion — that is, the Uving reefs are merely growing on 
parts of a submarine plateau where conditions favor their life. 

i Hayes, C. W.. Physiography and geology of region adjacent to the Nicaragua Canal route, Geol. See 
Amer. Bull., vol. 10, pp. 285-348, pis. 30-32, 1890. 
s Idem, p. 348. 

* Amer. Geog. Soc. Boll., vol. 46, p. 429, 1914. 

* Wash. Acad. Sd. Joom., vol. 6, pp. 57, 62, 1016. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 301 

The shore-line phenomena of Panama and Costa Rica have been 
carefully described by D. F. MacDonald in his forthconung report 
on the physiography and geology of the Canal Zone and adjacent 
areas. His condnsions.in general accord with those I have expressed 
for other areas. 

SOICE OTHER WEST nn>IAN ISLANDS. 

R. T. Hill in 1899 ^ pointed out '' that Jamaica was once a more 
extensive land than now, with benched and terraced margins which 
were submerged by subsidence/' and that ''similar submerged 
plains are now occupied by the growing reefs aroimd the island." 
Hill appears to hold the view that the reefs were formed during 
uplift, after submergence, and as regards the elevated fringing reefs 
I believe he is correct. In fact, Mr. Meinzer and I make a similar 
interpretation of the conditions imder which the coral-reef terraces 
of Cuba were formed. But, it seems to me that the barrier reef oflf 
Morant Point, Jamaica, has been formed after an episode of sub- 
mergence. The pouch-shaped harbors of Jamaica suggest that 
considerable stretches of the Jamaica shore line have undergone 
recent submergence. 

I have compiled information on the shore lines of other West 
Indian islands, but to present more seems unnecessary. Possibly 
except a reef off the southeast side of Barbados, all the off-shore 
West Indian reefs on which I have obtained information have 
formed on preexisting flats or plateaus during or after an episode of 
submergence. 

BRAZIL AND ARGENTINA. 

Herbert M. Smith,' it seems, was the first to recognize evidence of 
submei^ence on the east coast of South America, and Rich ' has 
made a pertinent apphcation of Smith's observations and deductions 
to the coral-reef problem. Smith says: 

Such an eetuary as I have described could only have been formed by the subsidence 
ol the land over a great area, and the encroachment of the sea on some former Amazons 
and its tributaries. 

During late geologic time there is in the region of the Amazon 
evidence of a higher followed by a lower stand on the land. 

Brannei^ has made the most careful study of the shore line of 
Brazil, and summarizes his conclusions as follows: ^ 

8. Although no changes of level are known to have taken place within the historic 
period, there are evidences of both elevation and depression of the Brazilian coast in 
late Geologic times. 

I Mas. Comp. ZooL Boll., toI. », pp. 99, 100. 

1 Smith, Herbert If., Notes on the physical geogrephy of the Amason Valley, Amer. NatunUst, vol. 
19, pp. 27-87, 1885. 

• Rkh, John L., The physiogiaphy of the lower Amawfi Valley as eyldenoe bearing on the ooralnreef 
problem, Sdenoe, new ser., vol. 45, pp. 689-990, June 8, 1917. 

i Branner, John Casper, The stone reefs of Brasll, their geological and geographical relations, with a 
efaapter on the ooral ntU, Mas. Comp. ZooL BolL, yol. 44, pp. 168, 189, 1904. 



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802 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

9. The evidences of depresBion coiisiBt of: 

(a) The open bays: Rio de Janeiro and Bahia. 

(b) The partly choked-up bays, such as Santos and Victoria. 

(c) The coast lakes formed by the closing of the mouths of estuaries such as Lagoa 
Manguaba, Lagoa d<( Norte, Jiquia, Sinimbu, etc. 

(d) Embayments altogether fiUed up. 

(e) The islands along the coast are nearly all close in shore and have the appeanance 
of having been formed by depression of the land. 

(/) Hie buried rock channels at Parahyba, now filled with mangrove swamps and 
mud, show a depression of at least twelve metres since those channels were cut. 
(g) Wind-bedded sand below tide level on Fernando de Noronba. 

10. The evidences of elevation consist of : 

(a) Elevated sea beaches especially well shown about the Bay of Bahia, and along 
the coast of the State of Bahia. 

(5) Marine terraces about Ilheos in the State of Bahia. These are about eight 
metres above tide level. 

(c) Horizontal lines of disintegration about one metre above high tide in granites 
and gneisses at and about Victoria, State of Espirito Santo. 

((f) Burrows of sea urchins so far above low tide that sea iirchins can not now live 
in them. These are well shown at Pedras Pretas on the coast of Pemambuco. 

11. Of the two movements the depression has been much the greater and was the 
earlier. 

12. The great depression probably took place in early Pliocene times. 

Additional evidence in support of the submergence of the Brazilian 
coast is given by O. P. Jenkins.* 

That the last dominant shift in the position of the strand line in 
eastern Brazil was by submergence, it seems to me, is incontrovertible, 
and that the Brazilian reefs are merely growing on the surface of a 
submerged continental shelf is too obvious to need defense. In these 
relations the Brazilian reefs accord with all other American offshore 
reefs, perhaps with the exception of the Barbadian reef specially 
mentioned on page 301. Professor Branner dates the submergence 
whereby the Brazilian harbors were brought into being, as Pliocene; 
whereas the submergence in the other areas discussed is clearly 
Recent. Without definite evidence I should not be justified in giving 
the drowning a later date than that assigned to it by lSx>fe68or Branner; 
but I now know that I assigned too great antiquity to some physio- 
graphic features I considered about the same time that he was engaged 
on his work on the Brazilian stone reefs; for instance, the higher Cuban 
terraces are Pleistocene and not Pliocene, as I said in the Cuba report 
previously cited. May not the antiquity of the submergence of the 
Brazilian coast be less than Professor Branner inferred? May not 
both the submergence and the minor uplift following it be post- 
Pleistocene in age ? Should the two events mentioned be geologically 
Recent, the shore-line history of Brazil would parallel that of eastern 
Central America. 

i JenUiis, O. P., GMlogy of the region aboat Natali Rio Grande do Norte, Bnan, Amer. Pfafios. Soe. 
Proc., vol. 53, pp. 431-406, 1913. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 303 

Willis has directed attention to two areas of submergence by down.- 
-warping along the Argentine coast, namely, the embayment of the 
Rio de la Plata and Bahia Blanca;^ but Barrell is of the opmioU; from 
the character of the submarine profiles, that there has been sub- 
mergence of the coast subsequent to the warping.' That there has 
been in late geologic time a rising of ocean level on the Argentine 
coast seems a justified deduction. 

ATLANTIC COAST OF THE UNITED STATES NORTH OF FLORIDA. 

That the last shift in position of strand line from the Georgia- 
Florida line at least to Narraganset Bay has been by submergence is 
so clearly shown by drowned stream mouths, resulting in estuaries 
and harbors, is so well known to geologists that no detailed presenta- 
tion of evidence is necessary. Northward from nesx Boston there 
has been subsequent to submergence, emergence, probably due to 
crustal reboimd after d^laciation and reUef of the pressure exerted 
by the superincumbent continental glaciers. 

TYPES OF WEST INDIAN AND CENTRAL AMERICAN LITTORAL AND 8UBLITT0RAL 
PROFILES AND THEIR RELATIONS TO CORAL REEFS. 

In my paper on littoral and sublittoral physiographic features of 
the Virgin and northern Leeward islands,' I pointed out that there 
are four types of subUttoral profiles in the West Indies (see fig. 19) 
as follows: (1) That found off volcanic islands, such as Saba, into the 
sides of which the sea has cut relatively narrow platforms; (2) fault 
plane profiles, such as the north side of St. Croix; (3) wide imdersea 
flats, where planation agencies have long been active, as off Anguilla 
and north of St. Thomas; (4) submarine banks, such as Saba, Pedro, 
and Rosalind, which have no bordering land, and whose upper sur- 
faces lie between 9 and 30 fathoms below sea level. All of these 
. areas have undergone geologically Recent submergence. 'Where 
do the offshore reefs occur ? 

There is no barrier reef on the fault slope on the north side of 
St. Croix. No reef started as a fringing reef, then increased in thick- 
ness and grew seaward so as to form a prism of coral-reef rock 
and material caught behind the reef, so as to become converted 
according to the Darwinian hypothesis into a barrier reef; but tnere 
is a barrier off the south side of the island, where gently dipping lime- 
stones pass below the sea and produce a platform on the surface of 
which at the proper depth a barrier reef has formed. Off the fault 
shore of the south side of Oriente province, Cuba, there is no barrier 
reef, but farther west, between Cape Cruz and Trinidad where there 

I wOUs, Bailey, a«ologlo notes, In Hrdllcka, A., Early man in South America, Bur. Amer. Bthn. Bull. 
02, I^ 16-18, 1912. 

> Barrell, Joseph, Factors in movements of the strand line and their results in the Pleistocene and post- 
PMstooeoe, Amer. Joucn. ScL, ser. 4, toI. 40, p. 6, IMS. 

• Washington Acad. Sol. JoanL^ vol. 6, pp. 68-M, 1916. 



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804 BULLETIN 103) UNITED STATES NATIONAL MUSEUM. 

is a submerged flat underlain by gently dipping limestones there are 
offshore reefs, some of which have the barrier form. Where there 
are extensive offshore flats at the proper depths, if the other eoologic 
conditions are favorable, reef corals grow upon th^ surface of the 
flats and form either patches, stacks, or barriers. 





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NORTH SIDE OF ST THOMAS 
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ROSAUNO BANK FUNAFUH ATOU. 

FIQ. 19.— TTPSS OP WB8T Iin>IAN SXXBUTTOBAL PBOFXLB8 AKD FBOIILB OF rVNATUTI ATOLL. 

It seems that no one woidd try to explain Saba, Rosalind, or Pedro 
Bank as the result of infilling behind barrier reefs. They are subma- 
rine plateaus, leveled by planation agencies, which almost certainly 
were both subaerial and submarine, and (hey have been submeiged in 
Recent geologic time. There is a rather meagre growth of reef corals 
on their windward sides; but these banks are scientifically of great 
importance, for, except that the coral growth is not so luxuriant, 
they essentially duplicate the great atolls in the Pacific. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 305 

SUBMERGED BANKS NORTH OF THE CORAL REEF ZONE IN THE WESTERN ATLANTIC 

-OCEAN. 

That there are otf the Atlantic coast of Central and North America, 
north of the temperature zone in which coral reefs now exist, sub- 
marine banks at suitable depth below sea level for the growth of reef- 
forming corals, has been stated in several of my papers.^ There are 
six submarine banks projecting seaward from the eastern part of 
Central and North America. Named in order from the south north- 
^irard these banks are, first, three on which there are coral reefs, 
namely, Mosquito Bank off Nicaragua and Honduras, Campeche 
Sank off Yucatan, and the Floridian Plateau; and, second, three on 
-which there are no coral reefs, namely, (jeorges Bank, the banks off the 
coast of Nova Scotia, and the Grand Banks of Newfoimdland. The 
presence of such banks is entirely independent of corals, but corals 
will grow on the surface of such banks where the necessary ecologic 
conditions prevail. 

SUMMARY OF THE CONDITIONS UNDER WHICH THE AMERICAN FOSSIL AND LIVING 

CORAL REEFS FORMED. 

1. The elevated Pleistocene fringing reefs of the West Indies are 
separated by erosion imconformities at their bases from the geologic 
formations that they overlie, but they were usually, if not invariably, 
formed during intermittent uplift following considerable depression. 

2. The offshore reefs, whether forming parts of more or less bedded 
formations or forming patches, stacks, or barriei^ of Uving reef, were 
formed during or after submergence, as is shown in the case of the 
fossil reefs by unconformable basal contacts wherever basal contacts 
could be studied, and in the case of the living reefs by a great variety 
of evidence indicating geologically Recent submergence. 

3. The offshore reefs grew upon or are growing upon antecedent 
fiUtts, 6nly a small part of the surface of which was or is covered by 
reefs. Ilie flats existed prior to the submergence during or after 
which the reefs developed. Corals are constructional geologic agents 
and help build up the sea bottom, but the large flats on which they 
grow would exist were there no corals. Such flats are not confined 
to the temperature zone in which* corals hve. 

4. The submergence of the basements of the fossil reefs seems more 
reasonably explained as the result of differential crustal movement; 
but the development of the Uving reefs seems in large part a result 
of geologically Becent rise in the stand of ocean level, for nearly the 
entire eastern shore of the Americas from Argentina on the south to 
Cape Cod on the north exhibits evidence of Recent submergence, after 
which there has been in some places minor emergence by differential 
crustal movement. The amount of the submergence usually seems 

> ScisDce, newter., vol. 41, pp. 508, 609, Apxil 2, 1916; G«ol. Soc. Amer. Boll., vol. 20, pp. 69-00, 1916; Amer 
Joani. Sd., ser. 4, vol. 41, p. 134, 1010; Carnegie Inst. Washington Yearbook No. 14, p. 238, 1910. 



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306 BULXETIN 1(JS, UNITED STATES NATIONAL MUSEUM. 

to be about 20 fathoms, but in places some facts indicate that the 
maximum is between 30 and 40 fatlioms. Although more accurate 
investigations of the amount of the submergence are needed, the 
available evidence accords with the hypothesis that glacial control i& 
one of the important factors in bringing about the formation of 
Uving coral reefs. 

GOBAL REBFS OF THE PaCIFIO OoBAN. 

It is manifestly impracticable to consider in this chapter more than 
a few of the important reefs of the Pacific Ocean. Those selected 
for discussion are the Oreat Barrier of Australia, the barrier r^f off 
New Caledonia, and those off the Fiji and Society islands. Finally 
a few paragraphs will be devoted to atolls. 

GREAT BARRIER REEF OF AUSTRALIA. 

The literature on the Great Barrier Reef is very extensive, and 
includes (Contributions from numbers of investigators, among whom 
Jukes, Saville-Kent, H. B. Guppy, Alexander Agassiz, A. C. Haddon, 
Wood Jones, E. C. Andrews, C. Hedley and Griffith Taylor, Eldge- 
worth David, W. M. Davis, and A. G. Mayer may be mentioned. 
R. A. Daly and I have based statements regarding it upon carto- 
graphic studies. No attempt will here be made to review all the 
literature, and attention will be mostly confined to those papers that, 
in my opinion, correctly interpret the relations of the reef. 

Andrews in 1902 published a remarkable paper ^ oh the shore line 
of Queensland and the platform on which the Great Barrier Reef 
stands. This paper contains an excellent account of the physiog- 
raphy of the Queensland coast, applying the deductions based upon 
the physiographic study to the conditions under which the reef 
developed, and in it is recognized the significance of a continuous 
platform and an interrupted reef. Because of the embayed shore 
line Andrews correctly inferred submergence of the AustraUan con- 
tinental shelf, and he makes the important statement: 

* * * the continuance in width of the shelf southwards of the limits of reefe 
(coralline), and the great shoals thereon, points to a minor part only of the shelf being 
formed of coral growth.' 

A few years later Hedley and Griffith Taylor published .a valuable 
paper on the same subject.' They accepted Andrews's deduction 

I Andrews, E. C, Prelininary note on the geology of the Qoeensland coast with referanoes to the 
geography of the Queensland and N. 8. Wales Plateau, Linn. Soc. N. 8. Wales, Proc for 1902, pt. % pp. 
14&-186, 1902. 

s Idem, p. 177. 

• Hedley, 0., and Taylor, T. Griffith, Coral reefs of the Great Hairier, Queensland: A study of their 
stmoture, life distribution, and relation to mainland physiography, Australasian Assoc. Adv. ScL, Ade> 
laide Meeting, Jan. 1907, pp. 894-418, 3 pis. 1908. 



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GEOLOGY AND PALEONTOLOGY OF THE OANAL ZONE. 



807 



regarding submergence and devoted particular attention to the 
effects of wind-induced currents in shaping atolls. They also say: 

It may be allowed, though Darwin deprecated the idea, that the continental shelf 
ipras ready prepared with numerous banks representing eroded islands, just reaching 
to \ritliin the required distance of the surface, when the first coral builders came. ' 

On a subsequent page they add: 

Whatever the history of the Great Barrier Reef was, the reefa of the Coral Sea, such 
aa Liihou Reefe, Flinders Reefe, and Herald Cays, shared in it. ' 

I have stated in one of my papers :* 

An inspection of the admiralty charts for the eastern coast of Australia shows con- 
clusively that the platform on which the Great Barrier Reef of Australia stands ha9 





9^>» nHftt- 



FlO. 30. — PSOmSS ACBOeS OOMTINKKTAL SHXLT, vast 8IDK OF AVSTEALU. THI LATITUPB at THS nfTEB- 
SBCnON OP KACH PBOPILB WITH THX 8H0BX UXfB IS POLLOWSD BT A SZATBMBKT OP THX DIBKCnON OP 
TBS PROnLE FBOM THE SHOBE. 
SOXTTH OP THE 80UTHEBN Ein> OP THE OBEAT BABEIEB REEP: 

1. FBOM SHOBE EAST OP LEADINQ HILL, 8. LAT. 25* 20' 15'% SOUTH 82* EAST. 

2. FaoM BASE OP Sandy Cape, S. Lat. 24*, 53' 40", Nobth 68* East. 

3. FBOM ToowoNO Hnx, S. Lat. 24* 22' 4", Nobth 45* East, passino between Ladt Eluot and 

lradt musobove islands. 
Across the Qbeat Babbibb Keep: 

4. FBOM RoDD Peninsula, 8. Lat. 24* (K 0", Nobth 50* East. 

5. Fbom Geoboes Point, Hincbinbbook Island, 8. Lat. 18* 25' 40", Nobth 72* SO' East. 

an existence independent of the Great Barrier Reef, and that corals have established 
themselves on this platform where the conditions favorable for their life are realized. 

Daly has given cross-sections of the AustraUan shelf both south of 
and across the Great Barrier Beef in two of his papers,^ and I have 
presented a series of cross-sections in one of mine,^ both of us basing 
our profiles on the British Admiralty charts. There is one important 
fact shown by both Daly's and my profiles, but which Daly seems 
not to have emphasized. It is that the platform not only continues 

1 Coral leefs o( the Great Barrier, Queensland, p. 406. 
t Idem., p. 413. 

« Washingtan Aoad. Sol, Joom., vol. 4, p. 32, 1914. 

i Daly, R. A., The gladal-^xmtrol theory of coral reefo, Amer. Acad. Arts and Sd., vol. 61, p. 107, flgi. 
21-24, 1015; Problems of the Padflo Islands, Amer. Joum. Sd., ser. 4, vol. 41, p. 170, figs. 26-20, Feb. 1016. 
6 Washington Acad. Sd. Joum., vol. 6, p. 64, profiles Nos. 1-^, 8-14, 1016. 



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308 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

southward from the reef limits, but in many places the barrier reef 
stands not on the margin of the shelf but miles landward from the 
edge. (See text-fig. 20.) There is also a significant terrace front 
at depths somewhat deeper than 120 feet. These profiles should 
be compared with those for the West Indies (fig. 11, p. 275). They 
tell essentially the same story. The platform can not be due to the 
presence of the Great Barrier Reef, for in many places it projects 
beyond the reef. I state in my paper cited:. 

The evidence in favor of a shore line between 25 and 30 fathoms below present sea 
level is strong, if not conclusive, and supports the deduction that the living barrier 
reef is growing on what was a land surface in Pleistocene time, an interpretation 
essentially that proposed by E. C. Andrews in 1902.^ 

NEW CALEDONIA. 

I have seen no good account of the coast of New Caledonia, off 
whose shores is one of the most important barriers known. According 
to P. Marshall, ^*'the northeast coast is practically straight, but many 
inlets that form excellent harbours penetrate the southwest coast.'' 
The chart shows indentations in the north coast, although they are 
not so deep as those on the south. I find references to the shore-line 
features in two of Professor Davis's papers," and from them certain 
information may be obtained. The shore line is embayed, there 
are deltas mostly contained in the embayments between headlands 
that are strongly clifled on the sea front. The present barrier reef 
has developed subsequent to the truncation of the headlands and 
subsequent to the submergence that has caused the embayment 
of the coast. Just how much of the platform surmounted by the 

1 W. M. Dayis has published since the manuscript of this paper went to press an article entitled: The 
Great Barrier Reef of Australia (Amer. Joum. Scl., vol. 44, pp. 33»^50, Nov., 1917), in which he oritidMS 
me and others because we have not "satisfiMtorily explained'* the origin of the form of "the continental 
mass." Among the statements of Professor Davis is " Vaughan's view is based on the physiogcaphic 
investigations of parts of the eastern coast of Australia by Andrews (1903); * * *", after he had intro- 
duced two quotations from my paper on the littoral and sublittoral ph3rsiographic features of the Virgin 
Islands, etc., as given in abstract (Amer. Oeolog. Soc. Bun., vol. 27, pp. 41-45, 1916). Profiessor Davis 
has drawn an erroneous deduction regarding my cartographic studies of the Great Barrier Reef. They 
could not have been based on Andrew's woriE, because Andrews neither published nor made comment 
on a series of profiles across the Australian platform, such as those I had prqiared. Furthermore, my 
emphasis of the fact, which it seems I was the first to point out— namely, that the present Great Barrier 
Reef in places stands some miles landward from the margin of the continental shelf— and my deduction 
therefrom, that the platform can not be attributed to infilling behind the reef, do not warrant the inference 
that ''Vaughan * • * has excluded coral-reef agencies from any imrt In forming the platform itself 

* * V I not only do not know how the Australian continental shelf was formed, but I do not know 
how any one of a number of hypothec can be tested. I, therefore, endeavored to confine my discussion 
to matters on which evidence is procurable, and said nothing regarding the origin of the platfor m. 
Professor Davis advances the hyi>othe8i8 that the platform on which the present Great Barrier is growing 
is a ''mature reef-plain", formed in a previous physiogr^thic cycle, and that it has been recently sub- 
merged. Whether reefs in past geologic time formed a rampart on the edge of the Australian omtinenta) 
shelf and a plain resulted from infllHitg behind the barrier can at present be neither jxroved nor dispro?ed 
and on this subject I have expressed no opinion. 

s" Oceania," Handb. regionalen Geolojgle, vol. 7, Abt. 2, p. 23, 1912. 

* Davis, W. M., Shaler Memorial study of coral reefs, Amer. Joum. 8d., ser. 4, vol 40, pp. 232, 233, 310 
243,245, 270, 1915; Problems associated with the study of corals. The Scientific Montiily, vol. 2, fig. 15 oir p 
25, p. 27, 1916. 



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GEOLOGY AND PALEONTOLOGY OP THE OANAL ZOKB. 309 

Caledonian barrier is due to the cut and fill process of marine 
planatdon at and below sea level during the cliffing of the 
promontories and to the sediment deposited in the sea, derived 
through the erosion of mature valleys, I can not say with cer- 
tainty, but that so much material deposited in the sea would 
imder the influence of waves and cmrents form a submarine plain 
is a warranted deduction; and as the barrier reef b crossed by 
gaps and is discontinuous at both the southeast and northwest ends, 
the deduction se^ms safe that it is superposed on a submerged plat- 
form of antecedent existence. 

FIJI ISLANDS. 

That the barrier reefs off the Fiji Islands have developed during 
or after submergence of their basements is obvious from an inspec- 
tion of the charts to anyone f amihar with the physiography of shore 
lines. The numei:ous reproductions of British Admiralty charts in 
A. Agassiz's volume on the Fiji Islands* is valuable and convenient 
for such a cartographic study. That the indentations of the shore line 
in the Fijis are due to the drowning of the lower parts of subaerially 
formed valleys has been pointed out by many geologists, the first 
of whom appears to have been Dana, who saysP 

There Ib, further, not merely probable but positive evidence of subsidence in the 
deep coast indentations of the high islands within the great barriers. . The long points 
and deep fiordlike bays are such as exist only where a land, after having been deeply 
gouged by erosion, has become half submerged. The author was led to appreciate 
this evidence when on the ascent of Mount Aoraion Tahiti, in September of 1839*. 
Sunk to any level above that of five hundred feet the erosion valleys of Tahiti would 
become deep bays, and above that of one thousand^feet, fiordlike bays, with the 
ridges spreading in the water like spider's legs; and this is a common feature of the 
islands and islets within the lagoons of barrier islands. The evidence of subsidence 
admits of no doubt. It makes the conclusion from the Gambler group positive; 
and equally so that for Raiatea and Bolabola represented on the charts in Darwin's 
-^ Coral Islands;" the Exploring Isles and others of the Fiji group; and that for islands, 
great and small, in the Louisade Archipelago and in other similar groups over the 
ocean. 

This statement was misinterpreted by Davis as being confirmation 
of Darwin's theory of coral reefs,* which, as is more than once pointed 
out in the present paper (see especially p. 249), carries with sub- 
mergence an hypothesis of platform building. Evidence of sub- 
sidence does not prove that the flat lying between a barrier reef 
and the shore has been formed by infiUing behind the barrier. 

Daly made a definite statement in 1910 in a list of ''maximum 
depths recorded for the drowned portion of these valleys," in which 

1 AfMBlx, Alexander, The Islands and Coral Beets of Fiji, Mas. Comp. Zool. Bull., vol. 38, pp. 1G7, 113 
plates, 1800. 

• Dana, J. D., Corals and ooral islands, ed. 3, pp. 378, 274, 1800. 

< Davis, W. M., Dana's oooflnnatioa of Darwin's theory of ooral reefs, Amer. Jonm. Sd., ser. 4, vol. 35, 
pp. 173-188, Feb. 1018. 



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310 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

he includes Mbengha and Moala of the Fiji group.* Subsequently 
Davis, in several of his papers, cited and others have similarly inter- 
preted the estuarine character of the lower ends of the valleys. 

Were the platforms on which the Fijian reefs stand, or which they 
margin, formed by infilling behind barriers or are the reefs merely 
superposed on antecedent platforms? In 1914 I published the 
foUowing statement: 

Having presented criteria for recognizing the relations of continental and large 
insular platforms supporting barrier reeib to the presence of the reefs, islands such as 
those in the Society and Fiji groups may be considered. ♦ ♦ ♦ A study of the 
charts of barrier reef islands, as Viti Levu, Fijis, and Tahiti, Society Islands, shows 
that the platforms are independent of the presence of reefs, and therefore the rela^ 
tions in these islands are similar to those indicated for barriers off continental ahoree, 
for here the reefs are also superimposed on platforms antedating their presence. 

Plate 7 of Agassiz's work on the Fiji Islands, already cited, shows 
the continuity of the platform northward and westward from Ovalau 
without any margining barrier reef. In my opinion these relations 
clearly show that the reef, where it is present, is merely superposed 
on an antecedent platform, and that the suggestion of Davis, that 
the ^itire platform is due to infilling behind a reef which in places 
has ceased to grow, is farfetched. 

Recently E. C. Andrews and W. G. Foye have published impor- 
tant papers on the Fijis. Andrews in his paper says: 

The Viti Levu salt water arms, th^efore, with their contained deltas, suggest the 
submergence of the Viti Levu coastal lowland in recent time, with the consequent 
drowning of the lower portions of the river courses. 

The island is girt with a Great Barrier Reef, several hundreds of miles in length, 
broken here and there by passages. The present Great Barrier Reef, which rises to 
the level of the sea, has-thus, in all probability, been built up by coral-reef arganisms 
upon the submerged lowlands of Viti Levu.' 

Andrews similarly interprets the conditions of development of the 
barrier reef oflf Vanua Levu. The interpretations advanced by 
Andrews essentially accords with mine; that is, the reefs are super- 
posed on a depressed platform that was previous to its submergence 
•a coastal lowland. 

Foye ' makes the following statement regarding Viti Levu: 

In general the present coral reefe are developing on platf<mns which originated 
during the deposition of the coastal series.* 

Regarding Vanua Levu he says: 

I visited only the eastern and central portions of Vanua Levu. The modem fringing 
reefe are here developing either along the shore line of recently submerged volcanic 
rocks or on coastal flats formed of the fine ash swept from the elevated hills of sub- 

1 Daly, R. A., Pleistooene gladation and the ooral reef problem, Amer. Xoum. Sd., ser. 4, toI. 30, p. 306, 
November, 1910. 

* Andrews, E. C, Relations of ooral reefs to crust movements In the Fiji Islands, Amer. Jonm. So., 
ier. 4, vol. 41, p. 138. 1916. 

* Foye, W. O., The geology of the Fiji Islands, Acad. Nat. Sd. Proc., vol. 3, pp. 306-810, April, 1017. 
« Idem. p. 306. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 311 

marine tuffo. The meet recent movements have been differential, and while uplift 
has taken place at the southeastern side of the island) subsidence has occurred to the 
east and north. The modem barrier reef occurs where subsidence has taken place 
^ther due to tilting ca Suiting during uplift.' 

Concerning the Lau Islands, he states: 

Within quite recent times the islands have subsided 50 to 90 feet and the modem 
coral reefs are developing on the eroded and submerged platforms.' 

One paragraph of Foye's condnsions is as follows: 

The data assembled by Daly and Vau^^ian convince the writer that Pleistocene 
platforms exist very generally throughout the coral seas. Yet while this Ib true, the 
platforms in Fiji are post-Pleistocene in their development. The writer was unable 
to discover any evidence of Pleistocene wave-cut platforms.' 

The second one of Foye's papers ^ contains the foUowing signifi- 
cant statement: 

There is another method by which atolls develop. The limestone iriands are rapidly 
eroded to sea level by atmospheric solution. Evidence of this process may be seen 
in the diminiflhing limestone masses within the lagoons of many of the Lau islands. 
By tidal scour and wave action platforms are developed slightly below sea level. 
Examples of such plaliorms may be seen about Fulanga and Ongea. It is significant, 
however, that most of these islands have lagoons 10 to 15 fathoms in depth. Such 
depths can not be ascribed to erosion, but must be the result of recent submer- 
gence. * ♦ * 

The information bearing on the Fijis may be summarized as fol- 
lows: 

1. The fringing reefs have unconformable badal contacts, as do 
those of the West Indies. 

2. The barrier reefs are superposed on antecedent platforms of 
diverse origin during or after submergence. 

3. The submergence is concomitant with, if not actually due to, 
differential crustal movement. 

4. In that they were formed during or after submergence and are 
superposed on antecedent platforms, the offshore reefs of the Fijis 
accord with all others, perhaps except a Barbadian reef, so far 
considered. 

SOCIETY ISLANDS. 

TAHITI. 

That Tahiti had undergone subsidence is impUed in statements by 
Dana,' the opcasional harbors being mentioned in two places in his 
book. W. M. Davis says:* 

The cli£f-rimmed island of Tahiti, the largest and youngest of the group, has suffered 
moderate subsidence after its cliffs were cut, but its bays are now nearly all filled 
with delta plains; hence a pause or stillstand has followed its latest sinking. 

1 The gtcAogy o( the Fiji laiuids, Acad. Nat. Sot Proo., p. 306, Apdl, 1017. 

* Idem, p. 300. 

* Idem, p. 300, 310. 

4 Foye, W. O., The geology of the Lau Ishmds, Amer. Joura. Sd., ser. 4, vol. 43, pp. 343-860, May, IoIt. 

* Corals and coral islands, ed. 3, pp. 140, ISS, 24«, 347, 1890. 

* Amer. Joarn. Sci., ser. 4, vol. 40, p. 271, 1015. 

37149— 19— Bull. 103 ^9 



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312 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

The condition of the reef between Taunoa Pass and Point Venus 
is interesting in this connection. Alexander Agassiz has giv^i a 
good description of this part of the reef and reef platform and has 
reproduced the British Admiralty chart of it.^ Agassiz says: 

Reef patches, the remnants of a former barrier reef, extend westward from YenuB 
Point parallel with the shore of Matavai Bay, forming the chain of Toa Tea reefs, but 
they are merely patches of Nullipores, with here and there diminutive coral heads 
which have taken no part in the building of these reefe. 

There is along the Toa Tea Reefs a great break in the continuity of 
the reef, but the platform continues, irrespective of the presence or 
absence of a margining barrier. The depths in Matavai Bay, 16 to 17 
fathoms, seem to be the maximum, are about the same as in Papiete 
Harbor, outside which there is a well-developed reef crossed by 
Papiete Pass. These reefs, also, seem to me to have grown up dis- 
connectedly on a submerged coastal flat. 

intfAf.f.TO ISLANDS or THB SOOITT GBOUP. 

Alexander Agassiz has described each of these islands in his coral 
reefs of the Tropical Pacific,' and P. Marshall has made the observa- 
tions and deductions recorded in the following quotation:' 

This reef marks the edge of the platfonn of marine erosion as described by Agassis, 
but the original maigin of the land before depression as described by Dannn and 
Dana. * ♦ ♦ 

It is evident that if the coral reef rises on the edge of a platform of marine erosion 
this very erosion would have worn the spurs back in such a way that they would 
terminate in steep cliffs. In no instance at Huaheine, Raiatea, or Tahiti that the 
author observed, did the spurs have an abrupt termination. The lower slopes ci the 
islands are in all cases notably less steep than the upper slopes. 

The deep inlets that intersect the coast line of Huaheine, Tahaa, and Raiatea are 
cleariy due to stream erosion. Prolonged marine action would have shallowed or 
filled them up or at least would have built up bars of coastal debris across the entrances. 

The author ia therefore strongly of opinion that the absence of cliffs at the termina- 
tion of radiating spiirs, the presence of deep water in the lagoon, and of far-reaching 
inlets, prove that marine erosion has not had any influence on the form of these islands 
at the present sea level. ♦ ♦ ♦ 

Finally the deep inlets appear to be drowned stream valleys and their nature 
strongly supports the belief that they have been subjected to an important movement 
of subsidence. 

Mehetia is interesting in that it is a young volcanic island, with a 
strongly diffed shore, a very narrow or no platform, and no coral 
reefs around it, only a few coral patches. That the other islands, 
Murea, Huaheine, Raiatea, Tahaa, Bora-Bora, and Maupiti have 
undergone geologically Recent submergence and that the barrier reefs 
have developed during or after submergence, can not be controverted. 
Is the reef flat due to marine planation and to terrigenous sediments 

1 AgaasU, Alezaiuler, The ooral iMb of the Tropical Padflc, lioi. Comp. Zool. Mem., toI. 28, pp. 1^- 
15^, pi. 900, 1903. 
s Idem, pp. 140, 141, 150-107. 
* Marshall, P., Oceania, Handb. reeionalen OeoloKie, voL 7, Abt. 2, p. 18, 1913. 



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GBOLOGY AND PALEOKTOLOOY OF THB OAKAL ZOKB* 813 

carried by the streams to the sea prior to the submergence after 
which the hving reefs have formed ? Unless sediment was deUvered 
to the sea so rapidly that a coastal plain pushed forward beyond the 
interstream divides as to protect them from attack by the sea, their 
seaward ends should have been cliffed, should the flat have been 
f onned in the manner suggested. What are the submarine profiles 
ojff the spur ends? Are there submerged cliffs at the divide tips? 
One of Agassiz's illustrations' represents a cliff of considerable height 
at one place on the shore of Maupiti. In my opinion suflSicient evi- 
dence is not available to establish how the reef flats of these islands 
were formed, and they may be made to accord with whatever theory 
of reef-flat formation an author may prefer. Should it ultimately 
be proved that these barrier reefs accord with the Darwinian hypothe- 




FXO. 21.— DUORAM TO SHOW HOW ▲ UNBAB IISBV LYING AGBOflS THE WIND IS FOBMKD INTO A HOB9X8HOS. 
(AnXS BXDLBT AND OBimTH TAYLOB.) 

siSy a fen^ instances in which that hypothesis applies will have been 

discovered. 

Atolls. 

There are two kinds of atolls: Those of the first kind rise above 
relatively shoal-water platforms, and are represented by the atolls of 
the Great Barrier Reef of AustraUa, those of the Floridian reef-tract, 
and the/aro« of the Maldives. That there was never any central land 
area for these atolls is perfectly obvious. Hedley and Grifl&th Taylor, 
in their paper, cited on pages 245, 251, have shown how the atolls 
along the Great Barrier have been shaped by the prevalent, mostly 
wind-induced, currents; and I have shown in my papers on the Mar- 
quesas and Tortugas atolls that precisely the same principles apply 
to them. The principles involved are illustrated by the accompany- 

1 The Coral Reefs of the Tropical Pacific, pi. 104, fig. 4. 



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314 BULLETIN lOSy* UNITED STATES NATIONAL MUSEUM. 

ing diagram (%. 21) , which is copied from Hedley and Griffith Taylor. 
Stanley Gardiner has given good descriptions of the faros of the Mal- 
dives.^ He says in a footnote on the page referred to: 

The technical tenn atoll is derived from the Maldivan atolu^ 8igiiif3riiig a province 
for governmental purpoees. Thore are 13 of these in the Maldives, and many cohobI 
of the islands on separate banks, most of which have distinct encircling series of reef 
reaching the surface. Many of the individiial reefs are themselves ring-shaped wi^ 
pools of water several fathoms deep in their centers. There are obvious disadvantages 
in using diminutives of the terms atoll and lagoon as applying to such. They are 
situated on shallow banks, and many are actually laiger than some of the isolated ring- 
shaped reefe of the Pacific, which arise separately in the deep basin of that ocean. I 
therefore propose to borrow further the Maldivan terms, faro and vehiy the former 
signifying such a small ring-shaped reef of an atoll or bank and the latter its central 
basin. I, further, following the Maldivan use of the term vduj apply it to deep pools 
even in the long, linear, circumscribing reefs of many of the banks, as I conceive that 
such pools have in all these reefs on banks the same mode of origin. 

On page 171 of the same work, Gardiner says: 

Each large reef on the bank is a separate entity that has grown up and pursued its 
history by itself, influenced it is true by the reefe in its vicinity but never directly 
connected with them. It is only now that the bank is at all approaching the condi- 
tion of the perfect atoll. Having seen how small faro may be formed from their 
earliest beginnings, we now see in North Mahloe the further fortune of such atolls, 
their joining together where possible to form long linear reefs with the loss perhaps of 
the whole inner part of their own reefe. 

The second kind of atolls more or less margin and more or lees 
completely encircle the flat summits of eminences rising from oceanic 
depths. The Darwinian explanation of the formation of such atoll 
rings is illustrated by figure 5, page 242, of this paper. Have these 
atolls formed in accordance with the postulates of the Darwinian 
hypothesis, or have more or less perfect rings developed on the 
edges of submarine flats, with or without submergence ? 

The origin of the first kind of atolls has been ascertained with so 
high degree of probability that it amounts to certainty. They have 
been formed on relatively shoal submarine flats, during or following 
submergence, and have been shaped by the prevalent currents. 
But a basement platform for the second kind of atolls can not be 
traced beyond the atoll limits, at least in our present state of knowl- 
edge. However, in case of atolls of an area so large as Rangiroa, in 
the Paumotus, for instance, the presumption is against their deriva- 
tion from barrier reefs according to the Darwinian hypothesis. 
They are too large, ^d, as Wharton long ago pointed out, their 
bottoms are too nearly level. If the Darwinian explanation were 
true, lagoon floors should be concave, more or less bowl shaped. 
That small, flat, summit areas may result from subaerial degradation 
and marine planation is known in many instances. That volcanic 

1 Gardiner, J. Stanley, The fauna and geography of the Maldlve and Loocadl^e Archipelagoes, toI. 1, 
pt. 2, p. 155, 1001-1903. 



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OEOLOOY AND PALEONTOLOGT 07 THE CANAL ZONB. 



315 



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316 BULLBTIK 103, UNITED STATES NATIONAL BCUSETTM. 

piles may be cut to wave base is known, and on page 311 of this paper 
Foye is quoted on a process by means of which reduction of limestone 
masses to sea level or slFghtly below sea level is accomplished. 

In this connection Salt Key Bank, which Ues between the Straits d 
Florida, Santaren Channel, and Nicholas Channel (text-fig. 22), is 
interesting, as' it is 61 nauticltl miles long by 37 nautical miles wide. 
Except a few marginal islets and elongate keys, it ranges between 
3i and 8 fathoms in depth. Alexander Agassiz visited and described 
this bank ^ and says that it is composed of eohan rock similar to the 

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Bahamas. The bank looks as if it were once a part of the Bahamas 
and was dissevered by faulting between it and the Bahamas. 
Whether that suggestion is or is not true, there is here a large level 
bank, obviously not formed according to the Darwinian hypothesis, 
that might serve an atoll foundation. Saba, Pedro, and Rosalind 
banks in the Caribbean Sea have been mentioned on pages 303, 304. 
Figures 23-25 illustrate them. 

It is not practicable to work out the geology of the f oimdations of 
the Paumotuan and the Maldive and Laccadive stolls, but the 

1 A reoonnaliwanoe of the Bahamas, etc., Mas. Comp. ZooL Ball., vol. 36, p. 81, pis. 1 and 31, 18M. 



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GEOLOGY AKD PALEONTOLOGY OF THE CANAL ZONE. 



817 



probability seems distinctly in favor of their being submerged pla- 
teau surfaces, upon which coral reefs, mostly marginal, have estab- 
lished themselves during and subsequent to moderate submergence. 
I will revert to Admiral Wharton's emphasis of the levelness of 
the floors of atoll lagoons (depth 24 to 26 fathoms), to his statement, 
''inside the low rim of growing coral which encircles their edges in 
various degrees,'' and to his question '^What causes this remarkable 
similarity of depth and this extraordinarily even surface over these 
large banks?" As 1 beUeve this short article by Admiral Wharton 
is one of the truly great contributions to oiu* knowledge of coral reefs, 




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FiQ. 24.— Chart o7 Fbdro bank. Fbom U. 8. rtdboobaphic chabt No. 1200. Scale 1 nrcH - 48 

NAUTICAL MILBS. 

the temptation to quote all of it is great. In it he points out one of 
the fundamental defects of the Darwinian hypothesis, namely, that 
the lagoon floor is not basin shaped as it should be if the atoll is due 
to the upgrowth of a reef that began on the slopes of a volcanic cone. 
He says: "I have no hesitation in saying that a flat floor is an 
invariable characteristic of a large atoll, and I can not find his 'deeply 
concave surface' in any large atoll. On the contrary, a flat surface 
is foimd in all of these, whether the rim be above or below the surface." 
Daly in his two papers dted has made an elaborate study of the 
depths of atoll lagoons of the Pacific and Indian oceans and has com- 
pared the depths in them with the depths in the lagoon channels of 
the same region. As the data compiled by him can not be repeated 



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318 



BULLETIN 103, UNITED STATES NATIONAL MUSEUIC. 



here, his later discussion in his paper on the Glacial-control theory 

may be consulted.^ Daly says: 

. ** Since probably not more than 5 m. to 25 m. can be allowed for the tMckneeB of 
the poBt-g^dal calcareous veneer in the wider lagoons, the accordance of platfonn 
deptn for the wider lagoons and reefless banks seems clear. Their range of 60-90 m. 
represents magnitudes of the same order as the depths computed for the Pleistocene 
wave-formed benches.** 

I have pointed out the similarity in the depths on Saba, Pedro, 
and Rosalind banks, to those on the atoU-lagoon floors of the Padfic. 
and Indian oceans — that is, the depths are between 20 and 30 fathoms. 




FlO. 36.— CHAST Of ROSALnrD and SABBAimXA BAinCS. FaOM U. 8. HTDEOOKAPmC CHAXT No. 964. 

The possibility of the formation of atoU lagoons by submarine 
solution was eliminated in the discussion on page 260 of this paper. 
Atoll rims are formed by constructional processes. That the greater 
abundance and luxuriance of reef-forming organisms on the periphe- 
ries of atolls is due mostly, if not solely, to the intolerance of such 
organisms to sediment, is shown by certain of my experiments. If 
the colonies are protected from sediment, the growth of corals within 
a lagoon may exceed that of corals on the outside. 

It is my beUef that the coral reefs forming atoU rims are superposed 
on platforms that antedate the formation of the living reefs, and 
which have undei^one a moderate submergence in Recent geologic 

» Amer. Acad. Arts and Sd., vol. 51, pp. 178-199, 1915- 



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QBOLOGY AND PAUBONTOLOGY OP THE CANAL ZONE. 319 

time. It is reasonable to ascribe this submergence to rise in ocean 
level because of deglaciation, because the order of magnitude of the 
submergence is the same as the order of magnitude expected from 
d^laciation. Mai^nal wave-cut benches should exist, or shguld 
have existed around the atoll banks. Perhaps more accurate hydro- 
graphic surveys and more detailed studies of the submarine profiles 
will discover them.. 

Conclusions. 

The results of an examination of the Tertiary, Pleistocene, and 
living coral reefs and reef corals of the West Indies, Central America, 
and the Southeastern United States are as follows: 

1. The fringing reefs have formed usually, if not invariably, during 
periods of intermittent uplift, following considerable submergence. 

2. All the important offshore reefs, both fossil and living, possibly 
except the reefs off the southeast coast of Barbados, have devel- 
oped during or following submergence after the subaerial erosion of 
their basements. 

3. Most of the fossil offshore reefs, all of those on which informa^ 
tion has been obtained, and all of these living reefs are superposed on 
antecedent flattish bas^nenta or platforms. Where there are no 
platforms, as off fault shore lines and young volcanic islands, there 
are no offshore reefs. 

4. Although corals are constructional geolc^c agents, they are 
subordinate to other limestone forming agencies, and none of the 
American platforms were formed by infilling behind a barrier. 

5. Submarine flats and plateaus at proper depths below sea levd 
to have furnished basements for offshore reefs are not confined to 
the temperature zone suitable for coral growth. Such extralimital 
banks are Oeorges Bank, the banks off the coast of Nova Scotia, and 
the Grand Banks of Newfoundland. Reefs form on such banks 
where the proper ecologic conditions for the life of reef building 
corals prevfdl. 

6. The submergences during and after which the fossil reefs were 
formed were almost certainly due to differential crustal movement; 
tjie submergence of the basement of the living reefs is probably due 
to complex causes, for there was differential crustal movement in 
the area under consideration dxiring Pleistocene time, also at some 
places within it during Recent time, and, in addition to these more 
or less local movements, there seems to have been during Recent 
time a general submergence of the eastern coast of America from 
Argentina to New England. The amount of the general Recent sub- 
mergence Ues between 40 and slightly more than 20 fathoms; an 
amoont of the order of magnitude that would be expected to result 
from the effect of deglaciation in raising sea level. The principal 
wave-formed Pleistocene plain now lies between 26 and 36 fathoms 



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820 BULLETIN 103, XJKITED STATES NATIONAL MUSBUIC. 

in depth, aad is separated by an escarpment from a shallower plain 
that now ranges between 17 and 20 fathoms in depth. What appear 
to be marginal hanging valleys north of St. Thomas and on the St 
Martin Plateau, and solution weUs, in the Bahamas, 33 to 38 fathoms 
deep, suggest that there may have been a short stand of sea level 
about 40 fathoms below its present stand. 

7. The fact that the terrace flat between 17 and 20 fathoms in 
depth is cut away on promontory tips on the windward side of St 
Thomas, while it is preserved in protected areas, indicates that the 
higher flat is older than the lower, and that it has been resubmerged 
aftw the development of the lower flat. The general similarity of 
the submarine profiles off Antigua, on the St. Martin Plateau, and on 
Mosquito Bank favors the inference that there was in those areas a 
similar lowering and subsequent rise of sea level. The subm^-ged 
channel within the channel at the mouth of Habana Harbor, and 
similar phenomena at other localities aroimd the Cuban coast, show 
that during later Pleistocene time Cuba stood more than 100 feet 
higher than nnmxediately previous to the cutting of these valleys 
within older valleys, and that after the valleys-within-valleys were 
formed there was submergence to an amount of about 100 feet. FaU 
of sea level during Pleistocene time and rise duiing Recent time is 
indicated for the Bermudas, the Bahamas, Florida, Central America, 
and the mouth of the Amazon, as well as for the areas just m«itioned. 
These phenomena are in essential accord with the demands of the 
Glacial-control hypothesis. 

8. The principal living West Indian and Central American reefs are 
superposed on submarine flats or plateaus of pre~Plei9tocene ago, that 
were dry-land areas during at least a part of Pleistoc^ie time, and 
while they were dry land they were wave cut and remodeled around 
their margins by submarine planation. 

9. There are two kinds of atolls, namely, (a) those that rise above 
relatively shoal-water platforms and were shaped by the prevalent 
currents, which are largely wind induced; (b) those that more or less 
completely encircle the flat summits of eminences that rise from 
ocean depths. These rings are formed by constructional geologic 
agencies, because, as submarine solution by sea water in such areas 
and at such depths is chemically impossible, a lower, flat area, sur- 
rounded by a higher rim can not be formed by submarine solution 
or by any other known destructional agencies. The depths on such 
banks as Saba, Pedro, Rosalind, etc., indicate that they were in 
large part, at least, above water during part of Pleistocene time, 
and that the flat simunits are largely due to processes operative in 
pre-PleiMocene time. What the processes were that caused the 
leveling of the sununits is a matter of pure speculation, but it seems 
probable that they were subaerial erosion and submarine planation. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL Z02^. 321 

The living coral rims on the banks enumerated have formed during 
and subsequent to Recent submergence. 

A review of the conditions imder which the principal barrier reefs 
in the Pacific Ocean were formed leads^ to essentially identical con- 
clusions. Those of the Australian Great Barrier, of New Cale- 
donia, the Fiji Islands, and Tahiti are superposed on antecedent 
platforms that have been submerged in Recent geologic time. The 
submergence of the Australian continental shelf apparently can be 
assigned to Recent rise of sea level because of deglaciation, as it 
seems that most of the surface of the platform was exposed as a dry- 
land area by withdrawal .of water from the ocean during at least a 
part of Pleistocene time. The submergence of the Fijian platforms 
is concomitant with, if not entirely due to, differential <srustal move- 
ment. The superposition of the barrier reefs off the shores of the 
smaller Society Islands on antecedent platforms is not proved. 
Evidence sufficient for the basis of an opinion is not available. The 
absence of reefs around Mehetia, where there is no shore platform, 
is significant. That the barriers off the other smaller islands were 
formed after the submergence of their basements is clear. The small 
cliffs at the spurs ends, in my opinion, do not constitute evidence 
against the presence of shore platforms, flats, or lowlands, ante- 
cedent to submergence. That ocean level in the Indo-Pacific, 
because of deglaciation, in Recent time has risen to an amount of 
about 60 meters (about 33 fathoms) as postulated by Humphreys 
and Daly, and that this rise of ocean level had - influenced the 
development of Uving coral reefs, is, I beUeve, so well established 
as to be almost if not quite incontrovertible. 

The rims of the large atolls, and perhaps of the smaller ones also, 
are growing, ui my opinion, on the surfaces of, mostly the edges of, 
flat smnmit areas that have undergone geologically Recent submerg- 
ence. These flats, I beUeve, were mostly formed in fre-Pleisto- 
eene time, and it is my opinion that they were largely put of water, 
or were very near the surface of the water, dmring Pleistocene time. 
If they projected above the water for an appreciable time, they should 
have been wave cut around their edges by the lowered Pleistocene 
sea, and evidence of such benching should be sought. I beheye the 
evidence will not be found on the hydrographic charts at present 
available, for the object of the published charts is to guide navigators 
rather than to serve as a basis for physiographic studies of the sea 
bottom ui depths where navigation is safe. 

From what precedes I believe it is clear that I consider that there 

' are two factors that determine the vigorous development of offshore 

reefs, which imder the most favorable conditions form barriers or 

atoll rims, the other proper ecologic conditions also being present. 

The first factor is the existence of an offshore flat, which may have 



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322 BULLETIN 103, UNITED STATES NATIONAL BCUSEUM. 

a land area on one side and open ocean on the other or which may 
be the top of an oceanic emineilce. The second factor is^gradual sub- 
mergeiiee. The vigor of offshore reefs where these conditions pre- 
vail can be correlated with certain ecologic demands of reef-forming 
corals. 

Reef corals thrive on offshore flats, near or against ocean water, 
because they are there removed from the deleterious effects 
of both land-derived and other sediment. Some of these rela- 
tions are well exemplified in the barrier reef off the east side of 
Andros Island, Bahamas. This reef grows on the outer, windward, 
edge of a small shallow flat, against the deep water of the Tongue of 
the Ocean. As the winds set landward across the reef no oceanic 
or land-derived sediment is deposited on the reef, it is bathed by 
the purest ocean water, and receives the largest amount of animal 
plankton that that part of the sea can supply. On the great shoals 
of the Bahama Banks and in the shoal waters of Florida behind the 
reefs the winds stir up the mud on the bottom ; the sediment while 
in suspension kills the plankton ; when it settles it kills those bottom- 
living organisms that can not endure being covered by mud. On such 
flats reef-forming corals can not live. On shallow banks coral 
reefs therefore thrive best on the windward sides. However, if the 
flat extends far enough offshore for land-derived sediment not to 
reach the reef and if the depth is sufficient for waves under ordinary 
conditions not to stir up the mud on the bottom, but not too deep 
for the growth* of reef corals, barriers may develop on the leeward 
sides of islands. A land area to the windward may actuaUy 
favor coral growth, as it breaks the force of the winds. A position 
on an offshore flat, particularly on the windward edge of a flat, 
insures a supply of the purest ocean water and an abundance of 
animal plankton. 

The gradual submergence of an offshore flat perpetuates the favor- 
able conditions for the life of reef-building corals, and gives an 
opportunity for continual growth upward. With upward growth 
during slow submergence of the basement the ecologic conditions 
for the life of reef-forming corals are made better, for the deleterious 
effects of sediment are minimized. 

As regards the life of corals, the method of bringing about these 
conditions is of no importance. . Whether the flat was formed by 
marine planation, by alluviation and the building of a coastal flat, 
by base-leveling through subaerial erosion, by the formation of a 
submarine plain of deposition, or by any other special process, is 
unimportant, provided the flat be formed. Whether the submerg- 
ence be caused by differential crustal movement, local or remote, or 
by rise in ocean level due to the melting of glaciers, is unim- 
portant, provided there be gradual submergence of the basement. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 323 

The maimer of producing the result is subordinate to the result. 
However the conditions may be brought about, preexistent flats 
and gradual submergence are two factors needed to supply continu- 
ously favorable conditions for the growth of reef-forming corals. 
The importance of deglaciation on modern coral-reef development 
consists in its having caused a gradual and moderate increase in the 
depth of the ocean, thereby producing submergence both in rate 
and amount favorable for the growth of reef-forming corals. 

The general conclusions here expressed are similar to those pre- 
viously published in a number of my papers. Before discussing the 
bearing of my conclusions regarding the formation of coral reefs on 
the theories advanced by others, I will give brief attention to some 
remarks by Prof. W. M. Davis. The following paragraph is copied 
from a paper by him entitled: The origin of coral reefs.* Similar 
remarks occur in others of his papers. 

Reefs and Reef-Flatfonns. A modification of Darwin's theory has lately been 
propoeed by Vaughan, who regards recent submergence proved by theembayments of 
the central islands as the determining cause for the upgrowth of existing biUrier reefs 
but who intei^rets the deeper and larger part of the entire reef mass as an independent 
"platform" of darlier origin. As this investigator has not yet published his views 
T^;arding the origin of the reef-platforms his modification of Darwin's theory will not 
be here discussed further than to note that it seems inapplicable to many barrier 
reefs in the Fiji and Society groups; that the discontinuity of certain barrier reefs 
eeems to be explicable on the assumption of imperfect upgrowth during and aftei 
a recent and rapid subsidence as well as on the assumption of independent origins for 
the reefs and their platforms; and that, while the extension of reef-platforms outside 
of the coral zone as in the case of the Great Barrier reef of Australia, truly suggests a 
dual origin of reef masses, this does not exclude the contemporaneous growth of plat- 
form and reef within the coral zone during long-continued but irregular or intermittent 
subsidence. 

Most of the objections raised by Professor Davis have been an- 
swered on preceding pages of this pai>er. It will be obvious to those 
who have read what I have said that my inferences as to submergence 
are by no means confined to the evidence of embajnnents in shore 
lines. In fact, many submerged areas show no clear-cut shore-line 
embajrments. It will also be obvious that the interpretation I am 
making did not originate with me. E. C. Andrews, in 1902, after his 
work on the Great Barrier reef of AustraUa, put forward in essential 
principles the same explanation. 

In answer to Professor Davis's statement ''regarding the origin of 
the reef platform,'* I will say that the recognition of the fact of super- 
position does not require knowledge of the constitution or origin of 
the basement on which an object or structure has been superposed. 
We may recognize the fact that a book lies on a table without knowing 
the kind of material of which the table is composed or the process of its 

1 Nat. Acad. SoL Proc., vol. 1, pp. 146-162, March, 1915. 



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324 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

manufacture; there is controversy as to the origin of the Sunderland 
terrace in Maryland and Virginia, but no geologist will deny that 
certain houses have been built on the surface of the Sunderland 
terrace flat; although the geologic history of the pre-Cambrian 
formations in Michigan and in other areas adjacent to the Great 
Lakes may be inadequately known, no one is justified in denying for 
such a reason that glacial deposits overlie the geologically old rocks, 
as it is obvious that the overlying material has in some way been 
placed on the underlying. The superposition of a geologic formation 
on another may be recognized without knowing the complete history 
of either the upper or the lower. The oligocene coral reef along 
Flint River near Bainbridge, Georgia, rests on the eroded surface of an 
upper Eocene limestone now designated the Ocala limestone. That 
knowledge of the Ocala limestone may not be adequate does not 
invaUdate the recognition of the facts that the fossil reef overlies it 
and that an erosion period intervened between its deposition and the 
growth of the reef, which obviously formed during or after the sub- 
mergence of its basement. 

To ascertain the origin of the submarine flats on which ofehore 
reefs stand is important in the advancement of our knowledge of 
geologic history, and I have acquired as much information on the 
subject as I could. I am completely convinced that there is no one 
explanation that can be applied to all of them. The following kinds 
have already been recognized: (1) Slightly tilted bedded tuflF, as in 
the fossil reefs of Antigua; (2) slightly tilted bed of limestone, as off 
the south coast of St. Croix and Cuba; (3) submerged coastal flats, 
as in the Fiji Islands; (4) submerged peneplained surfaces, as in the 
fossil reefs of Porto Rico; (5) submarine plains due to uplift of con- 
siderable areas of the ocean bottom and to the deposition of oi^anic 
deposits on such a siuf ace, as the Floridian Plateau previous to the 
formation of the middle and upper Oligocene reefs of Florida and 
southern Georgia; (6) flats of complex and not definitely known origin, 
such as those of the Antigua-Barbuda Bank, the Virgin Bank, and 
the continental shelves of tropical America and Australia. Plains 
suitable for the growth of corals have been formed by subaerial and 
submarine deposition, and by both subaerial base-leveling and sub- 
marine plantation. Nearly every, if not every, plain-producing 
process operative in tropical and subtropical regions' has taken part 
in the formation of plains on which corals have grown or are growing 
where the plains have been brought below sea level and where the 
other ecologic conditions for offshore reef formation obtain. 

I will revert to this subject in discussing the Glacial-control theory 
and in making suggestions as to future research. 



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GEOLOGY AKD PALEONTOLOGY OF THE CANAL ZONE. 326 

BbARINO Of THB8K CONCLUSIONS ON HtPOTHKSBS OF THE FoEkATION OF CORAL 

RXEFS. 

How do my results compare with the theories and hypotheses 
advanced hy others? Before considering my conclusions in their 
relation to those reached by other investigators, I wish to make a few 
general remarks on the literature appertaining to coral reefs. It is a 
subject that, in order to be properly treated, requires a considerable 
diversity of knowledge, as biologic, oceanographic, and geologic 
problems are involved. Very rarely has it been practicable for a man 
to be a specialist in all of these fields. Usually, as any investigator 
has been speciaDy qualified in only one or two of tbem, he has paid 
particular attention to those subjiBcts with which he was familiar, and 
nearly always did good work in those subjects; but in those fields in 
which he has been only casually engaged, his work is nearly always 
amateurish, and bis conclusions are in many instances erroneous. 
Should we expect a miLn who is primarily a biologist to be an expert 
in geology, edpecially when he attempts geologic work after he arrives 
at the place where he expects to conduct his investigations, without 
having had previous experience ? Should we expect a man who has 
riveted his attention on dry-land physiography, and who has not 
thought of biologic problems or of tha physiography of the sea 
bottom to take information from those branches of science? In 
reading the many publications on coral reefs, I am impressed with the 
particular, personal interests of the investigators, but what strikes 
me more forcibly is the excellence of nearly all the papers. I know 
no paper by a serious scientific man on a coral-reef area that does not 
contain records of valuable observations and correct conclusions. I 
have had the wish to write an accoimt of the very gradual growth of 
the knowledge we now have of coral reefs, and point out how each of 
the successive workers has contributed toward making that knowl- 
edge what it now is. It would be a record of honorable achievement. 
In the short review to follow I trust I may point out some of the 
substantial additions to be credited to those whose opinions I shall 
discuss. 

1 . The Darwin-Dana hypothesis, in my opinion, is correct as regards 
the formation of offshore reefs dming and after submergence; but as 
r^ards the formation of a prism of reef material, the upper surface 
of which forms a flat behind the barrier, their theory is wrong for 
every area on which we have definite information. Although the 
theoretic possibility of the conversion of a fringing reef into a barrier 
and a barrier into an atoll may not be denied, no instance of such 
conversion has yet been discovered. The inferences of Darwin as to 
areas of subsidence and of elevation, as shown on plate 3 of his work^ 
are largely in error, for barrier reefs are present where there is not 
general crustal subsidence, as Foye points out in his paper on the 



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326 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

geology of the Fiji Islands, wh^e ''since the Pleistocene period the 
algebraic sum of the movements has been positive and uplift has 
resulted.' ' * Very many similar instances, the Bermudas, the Bahamas, 
Florida, and Cuba among them, can be given. The criticisms of the 
Darwin-Dana hj^tiiesis apply to the recent publications of W. M. 
Davis. 

2. Semper, Alexander Agassiz, and others, who have maintained 
that barrier coral reefs have formed in areas of uplift, are correct, if 
the sum total of the movements since some date back in Tertiary 
time be considered, and their observations and deductions are valu- 
able in that they emphasize these facts; but they are in error in that 
they failed to take into account that in many areas there is incon- 
trovertible evidence showing submergence of the basements of the 
now-living reefs. Semper made astute observations on currents, but 
his deductions as to the formation of lagoons by destructional proc- 
esses are not warranted. 

3. Sir John Murray invented a very stimulating hypothesis, and 
correctly emphasized the necessity of taking submarine planation 
into accoimt in studies of the basements of coral reefs. He, however, 
overlooked important facts clearly proving Recent submergence in 
coral-reef areas, and his theory of the formation of atoll lagoons and 
lagoon channels through submarine solution by sea water is entirely 
disproved, and there are no other known destructional processes 
whereby lagoons may be formed, for lagoons are areas of sedimentation 
in which filling predominates over removal of material. 

4. Guppy is correct in his interpretation of oflEshore reefs being 
superposed on submarine platforms or "ledges/' and he made nu- 
merous valuable contributions to our knowledge of coral reefs, but he 
failed to take into accoimt evidence showing Recent submergence. 

5. Admiral Sir W. J. L. Wharton made one of the greatest con- 
tributions to our knowledge of atolls when he discovered the flatness 
of the floors and the imiformity of depth in atoll lagoons, and he 
pointed out the inadequacy of the Darwinian hypothesis to explain 
these phenomena. He emphasized the importance of submarine 
planation in leveling the top of peaks that reach or almost reach sea 
level, and definitely suggested the superposition of coral patches and 
atoll rims on flats produced in that way. He not only did not oppose 
the subsidence of such flats, but he thought that they frequently do 
*' subside and that some of the deeper lagoons may owe their depths of 
50 fathoms or so to such a movement, quite apart from subsidence 
of large areas which we know occurs." The only emendations of these 
statements that I can suggest is that the. probable effects of glacia- 
tion and deglaciation might have been considered. 

1 Nftt. Acad. Sd. Proc., voL 8, p. 809, 1017. 



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GBOLOGY. AND PALEONTOLOGY OF THE CANAL ZONE. 327 

6. Alexander Agassiz correctly observed the superpofiition of the 
living coral reefs of the Bermudas and the Bahamas on older lime- 
stone fomidations that stood above sea level previous to the sub- 
mergence which made possible the formation of reefs in the places 
where they now grow. He also pointed out the superposition of the 
Floridian, Cuban, and Central American living reefs on antecedent 
platforms or older limestone. He showed that in several areas in the 
Pacific the sum total of local crustal movements since some time in 
the Tertiary period had been upward. But he failed to take account 
of Recent submergence in Florida, the West Indies, and Central 
America, and he advanced the hypothesis that the living ofkhore 
reefs of the Pacific are superposed on wave-cut platforms without 
change of sea level by submergence of the land. I believe Agassiz 
correct in his emphasis of the need of an antecedent platform for the 
vigorous growth of offshore reefs; but he did not recognize the dear 
evidence of Recent submergence of the shores of the reef-encircled 
islands, and imfortunately tried to explain the formation of lagoons 
by submarine solution and scour. 

7. E. C. Andrews, I believe, is incontrovertibly correct in the 
essentials of his interpretation of the conditions under which the 
Great Barrier Reef of Australia has formed; that is, it is superposed 
on that part of the recently sid>merged Continental Shelf of Australia 
that lies within the temperature zone favorable for the life of reef- 
forming corals. 

8. Stanley Gardiner, who has made great contributions to our 
knowledge of Indo-Pacific corals and coral reefs and whose work on 
the. oceanography of the Indian Ocean is justly rated as classic, com- 
mitted the same errors in interpreting the geologic relations of coral 
reefs as did Murray and Agassiz. He failed to infer submergence 
from shore line characters and advocated the formation of lagoons 
through submarine solution by sea water. 

9. Hedley and Griffith Taylor agreed in all the essentials of 
Andrews's interpretation of the conditions imder which the Australian 
Great Barrier formed; they opposed Murray's solution hypothesis for 
the formation of lagoons, and correctly emphasized the importance of 
currents, largely wind induced, in the shaping of the atolls along the 
Great Barrier. 

10. Daly did not originate the Glacial-control theory of coral 
reefs, but he is its principal exponent. The following ascertained 
relations of living offshore coral reefs conform to the demand^ of this 
hypothesis: (a) They are superposed on antecedent basement fiats; 
(6) the amount of recent submergence, between 30 and slightly more 
than 20 fathoms, without deducting the amount of Recent up-build- 
ing of the sea bottom, which probably is as much as a few fathoms, 
is of the order of magnitude expected from deglaciation; (c) the 

37X40— 19— Bull. 103 ^10 



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328 BULLETIN 103, UNITED 8TATB8 NATIONAL MUSEUM. 

rate of growth of corals is known to be of such an order of magnitude 
as to account for the thickness of any known living coral reef by 
the growth of coral-reef organism since the disappearance of the 
last great continental glaciers. As Daly is not a specialist on corals, 
he has made some errors in his discussions of the geologic history 
and ecology of corals, but these errors do not affect the validity of 
glacial control being one of the dominant factors in modem coral^^ 
development. The only important point on which I am not in agree- 
ment with him is the evaluation of Pleistocene marine planation. I 
have shown that the Floridian Plateau has existed as a plateau at 
least since late Eocene time, and there have been extensive submarine 
flats in certain West Indian areas since late Eocene or Oligocene time. 
The submarine profiles that I have drawn for the West Indies, 
Central America, and AustraUa indicate Pleistocene benching in 
depths between 26 and 36 fathoms, without deducting anything for 
Recent upbuilding of the sea bottom. Certain West Indian and 
Central American reefs and the Australian Great Barrier, I, therefore, 
beUeve are growing on what were dry-land areas during at least a 
part of Pleistocene time. It, therefore, seems to me that many of 
the flats discussed by Daly are of pre-Pleistocene age, and that he has 
over-evaluated Pleistocene marine planation. Daly admits that there 
has been local crustal movement in some coral-reef areas. 

11. Wood Jones is undoubtedly correct in attaching great im- 
portance to the effects of sediment on the formation of coral reefs. 
No one who has had actual experience with coral reefs can for a 
moment doubt it. He also correctly accepts the interpretations 
of Andrews and of Hedley and Griffith Taylor for the Great Barrier 
of Australia, joining with the latter two in their opposition to the 
solution hypothesis and in their emphasis of the effects of wind- 
induced currents in shaping the segments of a reef. He, however, 
appears not to have appreciated the importance that, in my opinion, 
should be attached to submergence as factor in coral-reef formation. 

12. My own opinions can be very simply stated: (a) Fringing reefs 
seem always to have imconf ormable basal contacts ; they maybe formed 
after submergence that is not followed by uplift or during intermittent 
uplift that follows submergence ; that is, they may form during periods 
of either emergence or submergence of land areas. Are the basal con- 
tacts really significant ? Must not these contacts in the very nature of 
the case be unconformable ? If the basement has moved up with 
reference to sea level and a reef begins along the strand line, the 
basement of the reef will certainly be different from the reef itself 
and there will be an obvious unconformity. If the land mass sub- 
sides and a fringing forms along shore, the base of the reef wiU surely 
exhibit unconformable relations. I am unable to imagine a fringing 
without an unconformable basal contact. I never saw one that did 



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GEOLOGY AND PALEONTOLOGY OF THE OANAL ZONE. 329 

not have such a contact, (i) Offshore coral reefs, barriers, and 
atolls; form on antecedent flattish basements during and after sub- 
mergence in areas where the general ecologic conditions are suitable 
for coral growth, as stated on page 240. This generalization appUes 
to fossil as wt^ll as to living reefs, (e) Recent rise of sea level because 
of deglaciation has made conditions favorable for coral-reef formation 
over enormous areas, and it is one of the important factors in causing 
the great development of coral reefs at the present time. But in 
some areas, as in the Fijis, the flats on which the reefs are growing 
are coastal flats that have been brought below sea level by tilting, 
as described by Andrews and Foye. (d) The theoretic possibiUty of 
the progressive change of a fringing reef into a barrier and later into 
an atoll, according to the Darwin-Dana hypothesis, may not be denied, 
but no instance of such a transformation has as yet been discovered, 
(e) The coral-reef investigation is of value to geology, not so much 
because of what has been discovered regarding corals as it is that 
it has led to the study of a great complex of geologic phenomena 
among which corals and coral reefs are only incident. Further inves- 
tigations of the phenomena associated with coral reefs are among the 
great desiderata of geologic research. 

SUGOESnONS AS TO FUTURB InVESTIOATIONS. 

Before closing this discussion I will present a few suggestions that 
to me appear pertinent. 

1. It is my belief that, although ecologic notes are of much value 
in systematic work, not a great deal more advantage will result 
from such ecologic investigations in areas where corals are luxuriant 
as those conducted by Gardiner, Wood Jones, and others, including 
myself. We need to know more of the physiology of corals, but such 
researches must be conducted by expert physiologists. There is 
great need for ecologic work in the waters northward and southward 
from the coral-reef zone. Within the coral-reef zone there are three 
faunas delimited by depth and temperature. What happens outside 
the coral-reef zone? Do the deeper-water forms hve in shallower 
water as the high latitudes are attained ? Is it depth or temperature 
that causes the vertical faunal distribution within the Tropics? 
More knowledge of the ecologic relations of the deeper-water faunas 
in the Tropics and of the f aimas in both shoal and deep water in the 
temperate zones of the ocean is of great importance to geologists, 
for such knowledge would furnish a basis for interpreting the physical 
conditions under which some of the fossil faunas Uved. For some 
years I have wished to make an investigation of the kind outlined, 
but other duties have prevented the fulfilment of my desire. There 
is a large amount of morphologic work needed, both on the skeletons 
and on the soft parts of corals, but particxilar consideration of this 
subject is scarcely in place here. 



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330 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

2. The study of sediments in coral-reef areas has soarcely been initi- 
ated. Accurate determination of the source of the constituents of 
calcium-carbonate bottom-deposits should be made, the deposits should 
be classified according to their constituents, at least the area occupied 
by each kind of deposit should be ascertained as nearly as is practi- 
cable, and an endeavor should be made to ascertain the rates at 
which the different kinds of sediments accumulate. The results from 
investigations of this kind are of vital importance to geology, for 
only by firmly basing our inductions on wide and accurate knowledge 
of what is now happening in the ocean can we hope to make reliable 
deductions concerning the origin of and the conditions under which 
older sediments were formed. The quantitative evaluation of the 
work done by the different agents cooperative in the production of 
the different kinds of sediments should be an object constantly in 
mind. Although this is essentially a new field of research, during 
the past few years a number of investigators have notable achieve- 
ments to their credit. 

3 . Detailed studies of the general geology of tropical islands and con- 
tinental areas adjacent to tropical and subtropical waters should be 
imdertaken wherever possible. These investigations should include 
consideration of the stratigraphic and structural geology, the petrog- 
raphy of both the igneous and sedimentary rocks, very detailed work 
on the stratigraphic paleontology, and the physiognq>hy of the land 
areas. We now know that, by combining knowledge gleaned from the 
study of many relations, it is possible not only to recognize for an 
area the succession of rocks, their age equivalents in other areas, and 
their deformational history, but that it is also possible to ascertain 
the successive physiographic stages and other physical conditions 
throughout at least a considerable part of the history. The structural 
relations of the successive formations, the nature of the contacts of 
formations, and the character of the sediments, are among the criteria 
to be used in making the latter kind of deductions. Of how many 
tropical areas are there topographic maps on a scale of 1 : 62,500 or of 
1 : 125,000? Many areas, where the geology is very complicated, 
should be mapped on a scale of at least 1 : 20,006. The very detailed 
studies of a few carefully selected areas would supply keys for other 
areas and thereby accelerate work in other areas. Detailed woik 
of the kind su^ested should be done in Antigua, St. Bartholomew, 
St. Martin, and Anguilla, in the West Indies, for each of these islands 
typifies certain phenomena that are critical in elucidating the history 
of the West Indies, Central America, the southern United States, 
and northern South America. 

4. Biogeographic investigations supply a basis for deductions regard- 
ing former land connections and the dates of the separation of islands 
that may have been parts of large land masses. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 331 

5. Shore-line history is obviously an essential part of the study of 
coral reefs. But the entire story can not be deduced from the infor- 
mation furnished by all of the lines of investigation above suggested. 
The configuration of the sea bottom needs to be studied, both in 
plan and profile. Notwithstanding the great amount of work that 
has been done on oceanic hydrography, close attention to the minor 
configuration of the sea bottom and attempts to draw inferences from 
such studies are of very recent date. Since most hydrographic 
charts were not intended to serve as a basis for such researches, we 
are fortunate that we can extract so much information from them. 
Although it is probable that a much larger amount of data is on 
the charts than has as yet been utilized, that additional hydrographic 
research is needed is obvious. What are submarine slopes off the 
divide ends in reef-encircled islands) What is the character of the 
slopes off both the reefs and the breaks in the reefs ? The problem 
of submerged terraces, flats and fronts, has barely been touched. 
How extensively are such features present, and what is their signifi- 
cance) These considerations lead to inquiries regarding wave base, 
the rate of motion of the water, the erosional and transporting power 
of the water while in motion at different rates, and the relations of 
erosion and transportation to depth. Although the factors men- 
tioned are among those that determine the profile of subaqueous 
equihbrium and must be considered in their relation to it, there are 
other factors, among which are the initial slope of the bottom, the 
hardness and d^ree of consolidation of the material forming the 
bottom, and the attitude, height, and hardness of the rocks at the 
shore. More information on this complex of problems is urgently 
needed. 

Sea level rises or falls with reference to the land, or the land rises 
or falls with reference to the sea level. That there have been many 
shifts in the position of the strand line since the beginning of Pleisto- 
cene time is known to every geologist. He also knows that in many 
areas shifts have been caused by tilting or flexing of parts of the 
earth's crust, and that there must have been lowering of sea level 
while there were great continental ice sheets, followed by rise of 
sea level when the ice sheets melted. How much of the geologically 
Recent change in the position of strand line is to be attributed to 
climatic causes and how much to differential crustal movement? 
More accurate and areaUy more extensive studies of shore-Une his- 
tory should enable a more precise evaluation of the effects due to 
each than is now possible. Such investigations must not be confined 
to tropical and subtropical areas — they must be world wide. 

Then there is the problem of Pleistocene wave cutting. I beUev€>, 
for reasons stated elsewhere, that Daly has overevaluated the effects 
of Pleistoceme marine planation. Has either of us really enough 



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332 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

information to be convincing? Should answers to the questions 
raised in the preceding two paragraphs be forthcoming, and if we 
can make reliable estimates of the duration of the Pleistocene, the 
.amoimt of marine planation while sea level was lowered in the Pleis- 
^cene might be more nearly approximated. 

In conclusion, I wish to say that the questions and suggestions con- 
tained in the forgoing remarks have grown out of a study of corak 
^and coral reefs and the phenomena associated with them; and al- 
though it may have been shown, that corals are not so important as 
they were once considered to be, geolgists should be grateful for the 
romantic interest inspired by these lowly animals, for this interest 
has led us into the presence of some of the profoundest problems of 
geology. Perhaps the interest will endure and it may lead us to a 
better imderstanding of the world of which we form a part. 



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GBOLOGY AND PALEONTOLOGY OP THB CANAL ZONB. 388 
SYSTEMATIC ACCOUNT OF THE FAUNAS. 

Class ANTHOZOA. 

MADREPORARIA IMPERFORATA. 
FamUy SERIATOPORIDAE Milne Edwards and Haime. 

1849. Seriatoporidae ICilkb Edwards and Haime, Comptes Rend., vol. 29, p. 262. 

1869. PoeUloporidae Verrill, Essex Inst. Proc., vol. 6, p. 90. . 

1870. Stylophoridae Verrill, Conn. Acad. Arts and Sci. Trans., vol. 1, p. 514. 

In a recent publication ^ I have stated that while I seriously 
doubted the propriety of placing Stylop7u>ra and PodUopora in 
separate families, the traditional usage' was followed. Additional 
study since that statement was written has convinced me that Stylo- 
phora, Seriatopora, and PodUopora all belong to the same family. 
In fact, it seems that both Seriatopora and PodUopora are derived from 
StyJopliora, mostly through retrogression in the development of the 
septa. It is hoped to present in a future paper the evidence on which 
this suggestion is based. 

Genas STYLOPHORA Schweigger (emend. Milne Edwards and Haime). 

1819. Stylophara Schwbiooer (part), Beobacht. auf Naturf., pi. 5. 

1820. Stylophara Schwbiooer, Hand. Naturg., p. 413. 

1830. StylophofasjidSideropora db Blainville, Diet. Sci. nat., vol. 60, pp.319, 

361. 
1836. Anthopora Gray, Zool. Soc. London Proc. for 1836, pt. 3, p. 86. 
1846. Sideropora Dana, XJ. S. Expl. Exped. Zooph., p. 617. 

1860. Stylophara Milnb Edwards and Haime, Ann. Sci. nat., ser. 3, Zool., 

vol. 13, p. 102. 
1867. Stylophara Milnb Edwards and Hadcb, Hist. nat. Corall., vol. 2, p. 133. 

1861. Stylophara de Fromentel, Intr. Foljrp. foss., p. 179. 

1884. Stylophara Duncan, Linn. Soc. London Joum., Zool., vol. 18, p. 45. 

Thfpe-spedea. — Madrepora pisHUata Esper. 

Duncan in his papers on the Fossil Corals of the West Indies either 
describes as new or lists the following species: 
From the Ek)cene of Jamaica: 

Stylophora contorta (Leymerie) + 1 var. 
From the EJocene of St. Bartholomew, Cleve collection: 
Stylophora compressa * Dimcan. 
distans (Leymerie). 

1 Ctfnefie Inst. Washington Pub. 213, p. 73, 1918. 

• Attbough I hKV studied the collection rom 8t. Bartholomew submitted to Duncan, I could recognlie 
only one species whidi I have divided into our varieties. 



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334 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Styhphora eonferia Reuss. 

tvberom Reuss. 

# 

affimxB Duncan (described from Santo Domingo). 
grarwiata Duncan (described from Bowden, Ja- 
maica). 
From Santo Domingo: 

Styhphora qffinis Duncan. 

var. minor Duncan (a valid species). 
rarUteOa (Def ranee). 
From Bowden, Jamaica: 

Styhphora granulata Duncan. 
From St. Croix, Trinidad: 

Styhphora mimUa Duncan. 

raristetla (Def ranee). 

miraJnlia Duncan (not Duchassaing and Michelotti). 

I described in 1900 ^ Styhphora ponderoaa from the Oligocene of 

Salt Mountain, near Jackson, Alabama, and Styhphora ndnutissima 

from the Oligocene of Blue or Russell Spring, near Bainbridge, 

Georgia. 

I recognize as raUd the six species described as new by Duncan and 
the two later described by myself. Duncan's identifications of West 
Indian specimens with European species are all discarded as they are 
probably erroneous. 

In addition to the six species here described as new, I have de- 
scribed six other species in manuscript not yet published, making a 
total of at least 20 species of Styhphora known to me from the 
American Tertiary formations. The stratigraphic range of the genus 
in America is from the upper Ek)cene to Miocene. 



STTLOPHORA IMPERATORIS, mw 

Plate 74, figs. 1, la, 2, 3, 4, 4a, 5. 

Corallum attaining a rather large size, the basal part of some 
colonies as thick as a man's wrist. The cross-section of branches 
ranges in form from subelliptical to curved lamellate. The following 
are the diameters of the broken ends of the specimen, which is 62.5 
mm. long, represented by plate 74, figure 1. 

DiarneUrs in millimeter 8 of branches ofStylophora imperatoru. 



' 


Lesser 
diameter. 


Greater 
diameter. 


Basal eDd 






14.5 
13.0 
9.5 to 16 


— 
X7.0 




17.5 


Wider braDob 


34 










1 U. 8. Qeol. Survey Mon. 39, p. 133, 1900. 







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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 335 

The branch teiminals are compressed and often form sinuous 
plates. Thickness just below the summits about 3 mm.; width very 
variable, ranges from 6 or 7 up to 25 mm. Nodule-like growths are 
frequent on the sides of older branches. 

Cahces on older j>art8 of the corallmn from 1 to 1.3 mm. in diameter, 
therefdre rather large and conspicuous; intervening walls from 0.75 
to 2 mm. across, usually about 1.25 mm. Near and on the branch 
sunmiits the caUces are usually crowded and sUghtly less than 1 mm. 
in diameter. Upper margin of the caUces usually more prominent 
than the lower, sloping slightly downward, externally finely costulate. 

Septa, 6 primaries distinct, well developed, extending to the col- 
umella, the directives more prominent than the other primaries; 
secondaiies are small or obsolete, if they were present they usually 
have been destroyed in the type and paratypes of the species. 

Columella, a small, only slightly prominent style. 

Coenenchyma dense; its surface beset with pointed granulations. 

Localities and geologic occurrence. — Canal 2k>ne stations 6016, in ihe 
Emperador limestone, quarry. Empire, where some hundreds of 
specimens were obtained; 60246, lower end of culvert, Panama 
Railroad (relocated line), on Rio Agua Salud, in the upper bed, col- 
lected by T. W. Vaughan and D. F. MacDonald. Station 6026, in the 
Culebra formation, 2i miles south of Monte Lirio, Panama Railroad 
(relocated line), collected by T. W. Vaughan and D. F. MacDonald. 

Anguilla, station 6894, bli^, south side of Crocus Bay, in the lower 
50 feet of the exposure, collected by T. W. Vaughan. (See pi. 
74, figs. 4, 4a.) 

Doctor MacDonald obtained the specimen represented by plate 
74, figure 5, at station 1863 of the canal commission, on the west 
side of Gaillard Cut, between points opposite Cucaracha and Paraiso, 
station 5853 of the United States National Museum locality register. 
The specimen came from a layer, about 2) feet thick, consisting of 
pebbles, gravel, and tun's cemented with calcareous material; below 
the layer is gray, flaggy sandstone and tuff beds; above it is gray, 
fla^y sandstone, in thin layers separated by partings of carbonaceous 
black shale. The geologic horizon therefore seems to be in the Culebra 
formation, probably near its top. The specimen appears to be a form 
of StylopJuira imperatoris in which the caUces are more crowded than 
usual, as it agrees with that species in all other characters. 

Type.— No. 324752, U.S.N.M. 

Paratypes.— No^. 324753, 324754, U.S.N.M. 

STTLOPHOSA PANAMENSIS. Mw i 



Plate 75, figs. 1, la. 

Corallum, branches more or less contorted plates (see pi. 75, fig. 1). 
The thickness of the lower end of the type is 12.5 mm.; width, 
exceeds 28 mm.; length from base te summit, 38 mm. 



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336 BULLETIK 103, UNITED STATES NATIONAL MUSBXJM. 

Calicos small, apertures from 0.5 to 0.75 imn. in diameter; crowded, 
maximum distance apart 1 mm., usually less than 0.5 mm. — that is, 
less than a calicular diameter apart. Mai^gins very slightly or not 
at all elevated; upper wall in places forms an obscure upper lip. 

Septa, the six primaries distinct, fuse in the calicular axis, directive 
plane well marked; secondaries not recognizable in the type-speci- 
mens and appear to be absent, but it is possible that they were present 
and have been destroyed by fossilization. 

Columella a compressed style, not prominent. 

Coenenchyma, surface badly worn in the type, but some granula- 
tions may be distinguished. 

Locality and geologic occurrence. — Canal Zone, station 6016, in the 
Emperador limestone, quarry. Empire, collected by T. W. Vaughan 
and D. F. MacDonald. 

Type.—^o. 324763, U.S.N.M. 

S. panamensis has smaller and more crowded calices than S. 
imperatoris. 

STYLOPHOSA AFFINIS ] 



1863. Stylophora affinis Duncan, Geol. Soc. London Quart. Joom., vol. 19, p. 436, 
pi. 16, fig. 4. 

1866. Reuada affinU Duchassaino and Miohelotti, Sup. GoraU. Antilles, p. 70 
(of reprint). 

1867. Stylophora affinis Duncan, Geol. Soc. London Quart. Joum., vol. 24, p. 25. 
1870. Rettssia affinii Duchassaino, Rev. Zooph. Antilles, p. 26. 

Original description. — "Corallum branched, large; branches nearly 
cylindrical, leaving the stem at an acute angle, slightly flattened on 
one side. The largest stem is four-fifths inch in diameter. Blunt, 
aborted, branchlike swellings exist on some of the lai^er stems. 
Corallites radiating from the center of the stem and branches, sepa- 
rated by about their own width of dense coenenchyma, which is seen, 
in the larger specimens, to be very slightly cellular. Walls not distin- 
guishable from the coenenchyma in the substance of the mass, but 
slightly raised into a very shallow craterif orm edge on the surface. 
Calices circular, a very little raised as crateriform elevations, very 
numerous, disposed irregularly, but very nearly equidistant in some 
places and less so in others; margins sharp. Diameter one- thirtieth 
inch [0.83 mm.], rarely larger. The calicular margin, when well pre- 
served, looks like a little ring placed on the intercalicular space, and 
the small styliform columella renders the appearance very distinct. 
Intercalicular spaces marked by a continuous and rigid line, which, 
being in the part of the spaces at the base of the calicular eleva- 
tions, and being continued round each calice, is, from its general 
straightness, formed into irregular polygons. The line is sensibly 
raised, convex, and now and then dentated. Between the line and 
the calicular margin there are distinct papillae, one row at the very 



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QBOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 387 

mai^iiial edge, the other corresponding to it a little lower down the 
calicular wall; a third is sometimes seen; and in places where there 
is an tmusual distance between the calices, and when the 'line' is 
wanting, the papillae are numerous, distinct, and a little smaller. 
The line and the papillae form a very marked distinction. Between 
some calices there are faint elevations. Septa whole, not exsert, but 
little visible in perfect caUces, but very distinct when the coral is 
worn. Upper margin perfect and concave upward, the septa ap- 
pearing festooned to the columella; they are deUcate, very little 
thicker at the wall than elsewhere, and join the columella high up 
near its point. The papillae at the caUcular edge extend a little on 
the wall, and may be considered as rudimentary septa and costae; 
if so, there is a second cycle, and also a third in half of each system. 
The' persistence of six septa, nearly all of the same size, is very re- 
markable. Columella styliform, large and dense in the corallite, and 
forming a rotmded-off cylinder with a sharpish rounded tip, which is 
very distinct halfway down the calice. CaUcular fossa shallow, 
about half as deep as broad. Endothecal dissepiments stout, trans- 
verse, numerous. The walls and columella do not fill up the lower 
parts of the corallites. Increase by extracalicular gemmation. 

"From the Nivajfi shale. Coll. Geol. Soc." 

Duncan reports the species from the Nivaj6 and Cerro Gordo 
shales, Santo Domingo. 

I have received 22 specimens labeled Stylophora affinis from the 
Museum of Comparative Zoology, and 6 from the Philadelphia 
Academy of Sciences. I have separated four of the specimens be- 
longing to the former institution and have described them as a new 
species. Six specimens are S, affinis, 9 are worn but probably are 
S. ajfims, 2 seem to be different and possibly belong to a different 
species, 1 I refer to Duncan's S. grarmUUa. I think that two of 
Philadelphia Academy are referable to S, ajfvrm, the four others are 
probably worn specimens of the same species. 

In the specimens that I have referred to S. affinis the upper mai^n 
of the calice is more prominent than the lower forming a small, pro- 
jecting hp. Duncan's description in other respects is satisfactory. 
As the sTU-f ace of specimens is easily worn by rolling, the upper lip 
of the calice and the surface ornamentation being destroyed, the 
positive identification of many specimens is rendered impossible. 
On the tips of the branches, which are blunt and roimded, the calices 
are crowded, with no development of intervening coenenchyma. 
• Miss Maury obtained in Santo Domingo a single specimen, a piece 
of a small branch, of this species, on Rio Gurabo, zone D, associated 
with Madrads deeadis (Lyman), PociUopora crassoramosa Dimcan, 
Stephanocoenia intersepta (Esper), OrhiceUa limbata (Duncan), 
OrbiceUa cavernosa var. cylindrica (Dimcan), and SyzygopTiyUia 
dentaia (Duncan), 



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338 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

I collected at station 3446, in the La Cruz marl, first deep cutting 
east of La Cruz, near Santiago, Cuba, casts of the surface of a species 
of Siylophora. Squeezes of the surfaces of these casts agree com- 
pletely with specimens from Santo Domingo identifiidd by me as 
S, afflnis. I am therefore attaching that name to the specimens. 
It is probable that similar casts from other localities in Cuba repre- 
sent the same species. 



STYLOPHOBA PORTOBELLENSIS. mw i 

Plate 76, figs. I, la. 

Corallum ramose, branches compressed, more or less contorted 
flabellate at the terminals. Growth form, therefore, similar to that 
of Stylophora imperatoris. The type is 37.6 mm. long; smaller 
diameter of basal end 10 mm., width of base about 13 mm. ; maximum 
width of branch in horizontal plane about 22 mm., thickness at same 
level 10 mm. 

CaUces shallow, diameter averages about 0.75 mm. or sUghtly 
less; distance apart approximately equals the calicular diameter, 
in places less, 0.25 to 0.5 mm.; margins flush with the coenenchymal 
surface, in places slightly elevated on the upper side, but not enough 
to form a distinct upper lip. 

Septa, six primaries distinct, rather thin, extend to the columella; 
no vestige of secondaries was observed. 

Colum^ella, a pointed style, moderately prominent, thickened below 
the bottom of the cahce. 

Coenenchyma dense or costulate with an intercaUcular ridge and 
cells on its sides. The surface is worn, but vestiges of small granu- 
lations may be recognized. Axis of the corallum spongy. 

Locality and geologic occurrence. — ^Panama, probably from near 
Porto Bello, collected by D. St. Clair; geologic horizon imknown. 

Type.— No. 324762, U.S.N.M. 

This coral has considerable resemblance to StycopTiora goeOudsi, but 
its calices are distinctly largerj and their uppei* mai^ns are in some 
places slightly raised. Siylophora imperatoris has larger caUces with 
distinct upper hps. Stylophora portobeUensis appears most closely 
related to Stylophora affinis Duncan, from the Nivajd shale of Santo 
Domingo. 

6TTLOPHORA GOETHALSI, new apedes. 

Plate 76, figs. 2, 3, 4. 

Corallmn ramose, witJb branches subelliptical or much compressed 
in cross-section, in this character resembling S. imperatoris. Branch 
summits frequently or usually with digitiform protuberances (see 
pi. 76, fig. 2). 

Calices shallow, decidedly smaU, 0.5 to 0.75 nun. in diameter; and 
relatively distant, from a calicular diameter up to 1.5 mm. apart- 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 



339 



Calicular margins obscurely or not at all elevated; without a pro- 
tuberant upper lip. 

Septa, six distinct primaries, about equal in size, extend to the 
columella; secondaries much smaller, but can be distinguished in 
the better preserved caiices. 

Columella a small, slightly compressed, fairly prominent style. 

Coenenchymal surface closely set with pointed graaulations. 

Locality and geologic occwrrence. — Canal Zone, at stations 6016, 
quarry in the Emperador limestone, Empire, Canal Zone, collected 
by T. W. Vaughan and D. F. MacDonald; 6026, in the Culebra 
formation, 2\ miles south of Monte Tjirio, Panama Railroad (relocated 
line), collected by T. W. Vaughan and D. F. MacDonald. 

CoiypeB.—'^o. 324767, U.S.N.M. (3 specimens). 

Siylojihora goethcUsi resembles the Santo Domingan species, S, minor 
Duncan, whicii is ramose and has small caiices, from 0.5 to 0.75 nun. 
in diameter. The end of the branches in R. goefJudsi are more com- 
pressed than in 8. minor, its caiices are slightly larger, and its 
secondary septa are better developed. Although closely related, 
they appear to belong to distinct species. 

STYLOPHOSA MACDONALDI, new spedM. 

Plate 75, figs. 5, 5a, 6, 6a, 7, 7a. 

Corallum composed of elongate, slender, curved branches and 
branchlets, with bluntish, rounded summits. The only branch 
terminal that is perfect is represented by plate 75, figure 5. The 
following are measurements of four broken branches: 

Measurements in millvmeieTB of branches ofStylophora macdonaldi 



Branch No. 


Length. 


Diameter of 
smaller end. 


Diameter of 
larger end. 


1 


15.5 3. 5 by 5.0 
19.0 4.0 by 6.5 
21.5 4. 5 by 5.0 
28.0 4.0bv4.0 


5.0 by 5.3 


2 


5.5by7.5 


3 


5.0 by 6.5 


4 


4.5 by 5.6 











Just below the place of bifurcation. the parent branch is consider- 
ably compressed; in one branch the greater diameter below a fork 
is 1 2 mm., while the lesser diameter is only 6.5 mm. 

Caiices rather shallow, but distinctly excavated; diameter, 1 mm.; 
distance apart from 0.5 to 1.5 mm., usually less than the caUcular 
diameter; margins usually slightly or not at all raised, but knots 
correspond to the outer ends of the septa. There is no upper lip to 
the caiices. 

Septa, six well-developed, strong, subequal primaries extend to 
the columella; secondaries small but usually distinct. Subequal 
knots correspond to the outer ends of th^ two cycles of septa, and a 



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310 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

smaller knot with no corresponding septom usually occurs betweeo 
each pair of larger knots. 

Columella, a distinct, round, moderately prominent style, very 
slightly compressed in the directive plane. 

Coenencbymal surface roughly granulated, from 1 to 4 rows of 
granules between calices, depending on their distance apart. 

Localities and geologic occurreTice. — Canal Zone, in the Emperador 
limestone at stations, 6016, quarry, Empire; 6024ft, lower end of cul- 
vert, Panama Railroad (relocated line), on Rio Agua Salud in the 
upper bed, collected by T. W. Vaughan and D. F. MacDonald. 

Ootypes.—No. 324769, 324770, U.S.N.M. (7 specimens). 

Of other species of Stylophora with which I am acquainted S. mac- 
doruMi seems to resemble most S. granuUUa Duncan from Bowden, 
Jamaica. S. granuUUa has deeper calices, less developed secondary 
septa, and in some specimens the upper lip of the calices is nKHie 
prominent than the lower. 

STTLOPHOBA GSANULATA Dwmuu 

1864. Stylophora granulata Duncan, Geol. Soc. London Quart. Jour., vol. 21, 

p. 10, pi. 2, fig. 3. 
1867. Stylophora granulata Duncan, Geol. Soc. London Quart. Jour., voL 24, 

p. 26. 
1873. Stylophora granulata Duncan, GeoL Soc. London Quart. Jour., vol. 29, 

p. 551. 

Original description, — ''The corallum is ramose; the branches are 
nearly cylindrical, often flattened on one side, and leave the stem at 
an acute angle. The caUces are placed irregularly, and are separated 
by a coenenchyma, which is sharply granular, and which has veiy 
rarely any grooves or continuous ridges on its surface. The calices 
are circular, not inclined, very deep, and are surroimded by a raised 
ring formed by the septa and costae. The columella is situated 
deeply; it is cylindrical below, and sharp where free, but it does not 
reach the level of the calicular margin; it is delicate, and six luge 
septa are attached to it low down. The septa are in two sets. Hie 
superficial septa are from eighteen to twenty in number; six are con- 
tinuous with the large septa, and the rest taper finely internally and 
externally, the spindle-shaped process being one-half septum and the 
rest costa. The processes are close, radiate, and horizontaL Diam- 
eter of calices, one-thirtieth inch [0.8 mm.]. 

''Localities: Bowden and Vere, Jamaica.'* 

Duacan, in 1873, cites this species from St. Bartholomew, but this, 
I am convinced, is an erroneous identification. 

There are two small broken branches of this species in the collec- 
tion of Mr. T. H. Aldrich, obtained at Bowden, Jamaica, and pre- 
sented to the United States National Museum. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 341 

Specimen No. 1. — Small branch, 16 mm. long, diameter of lower 
end 4 mm.; upper end flattened, bifurcating, greater diameter 5.5 
mm., lesser 3 mm. 

Diameter of calices veiy slightly less than 1 mm., separated by 
about the same width of coenenchyma. The margin is usually a very 
slightly elevated rim without an elevated hp around which are 12 to 
18 small costae. In a few instances the costae continue from one 
calice to the next, but usually the intercalicular coenenchymal sur- 
face is merely granulate. There are from two to six indefinite zones 
or wavy lines of granulations between two calices. The granula- 
tions are subconical, round-pointed. Umits of zooids sometimes 
faintly indicated by a slightly raised granulated line. Calices mod- 
erately deep. Six principal septa, the second cycle represented by 
small short septa, variable number of rudimentary members of the 
third. The upper margins are sUghtly exsert. 

C!olumella does not reach to level of calicular mai^in, sharp- 
pointed. 

Specimen No, t. — ^A small somewhat compressed, broken branch, 
16 mim. long; greater diameter of lower end, 6.5 nmi., lesser, 5 mm.; 
greater diameter of upper end, 6 nam., of lesser, 4 mm. Diameter of 
calices very slightly more than 1 mm. Width of intervening coenen- 
chyma averages about the same as the diameter of the cahces. 
Calicular rim a little elevated, and slightly swollen around the base. 

Costae longer than in No. 1. Granulations about the same in both 
specimens. Elevated line between zooids usually distinct. 

There is in this collection a third specimen which is probably only 
a variation of the same species. It is a fragment of a branch 14 mm. 
long. The diameter of the caUces is about 0.75 mm.; the calicular 
rims are not elevated but usually tend to be depressed. The coenen- 
chymal surface is very densely and minutely granulate. The limits of 
adjoining zooids are indicated either by a very faint raised or by an 
impressed line. 

Localities and geologic occurrence. — ^Besides occurring in the Bowden 
marl of Jamaica, Stylophora granulata is also found in Cuba at sta- 
tions 3476, Baracoa, and 3461, gorge of Yimiuri River, Matanzas, 
collected by T. W. Vaughan. 

Santo Domingo, station 7781, Bio Cana, zone H, collected by Miss 
C. J. Mauiy. 

STTTLOPHOBA CANAUS. new spmIm. 

Plate 76, figs. 2, 2a. 

Corallum of type, a small, nodular mass, 42 mm. long, 23 mm. tall, 
and from 10 to 14 nmi. thick (see pi. 76, fig. 2, for view, natural size, 
of the upper surface). 



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342 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

CaUces shallow^ fairly large, 1 mm. in diameter; xisually 1 mm. 
apart. Margins not elevated; the walls barely distinguishable from 
the surroimding coenenchyma. 

Septa in two distinct cycles; only the six primaries readi ihe 
columella, but the secondaries are well developed. 

Columella, a pointed style. 

Coenenchymal surface crossed by costules, along which are rela- 
tively coarse granulations. In places the coenenchyma appears cel- 
lular, as the costules are not solidly fxised but have cellules developed 
between them. 

Locality and geologic ^occurrence. — Canal Zone, station 6016, in the 
Emperador limestone, quarry, Empire, collected by T. W. Vaughan 
and D. F. Macdonald. 

Type.— No. 324775, U.S.N.M. 

This species most closely resembles a species from the base of the 
Chattahoochee formation, on Flint River, 4^ miles below Bainbridge, 
Geoi^a, but it differs from the latter species in two characters, 
namely, the outer ends of the principal septa are not produced into 
prominent teeth, and in places the coenenchyma is distinctly cellular. 

STTLOPHOBA PONDEBOSA Vaagkuu 

1900. Stylophora ponderosa Vaughan, U. S. Geol. Survey Mon. 39, p. 132, pL 
13, fig. 16; pi. 14, figs. 1, la, 16. 

One of the specimens obtained by me in Antigua seems referable 
to this species. The upper surface has four nipple-shaped elevations 
on it; the largest is about 15 nmi. in diameter at the base, about 
5 mm. tall, and about 5 mm. in diameter just below the rounded 
summit. Except such protuberances, the surface is flattish, with 
some undulations. The size of the caUces and the septal characters 
are as in the cotypes of S. ponderosa. 

Localities and geologic occurrence. — ^Alabama, Salt Mountain, 6 
miles south of Jackson, just above the top of the Vicksburg group, 
collected by T. W. Vaughan. 

Antigua, station 6854, Rifle Butts, in the Antigua formationi 
collected by T, W. Vaughan. 

Genas POOILLOPORA Lamarck. 

1816. Pocillopora Lamarck, Hist. nat. Anim. sans Vert:, voL 2, p. 273. 
1918. PocUlopora Vaughan, Carnegie Inst. Washington Pub. 213, p. 75. 

Type species. — Pocillopora acuta Lamarck. 

Duncan desciHbed two fossil species of Pocillopora from the West 
Indies, P. crassoramosa * from the Nivajd shale of Santo Domingo, 
and Pocillopora tenuis ' from Antigua. I have seen good suites of 

1 Q«ol. Soc London Qnart. Jonrn., vol. 20, p. 40, pi. 6, figs. Qa, 9b, 1864. 
'Idem, vol. 24, p. 21, pi. 1, figs. 5a, 66, 6c, \wri. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 343 

specimens of P. crassoramosa, but have seen none of P. tenuis. 
P. crassoramosa has thickish branches on which verrucae may be 
wdl developed or obsolete; P. tenuis appears to be of more or less 
massive growth-form and has across the corallite cavities thiu 
tabulae, the spaces between which are not filled by steroplasmic 
deposit. 

I have specimens rei»*esenting four additional American, fossil 
species of the genus. They are all branching forms. I collected 
one of the species at Willoughby Bay, Antigua, in the Antigua forma- 
tion; and another in the upper Oligocene marl at Baracoa, Cuba. 
The specimen at the latter locaUty was obtained in association with 
Stylophora granulata Duncan, which was origmally described from the 
Bowden marl of Jamaica. Miss Carlotta J. Maury obtained P. 
crassoramoaa in Santo Domingo in what she designates z6ne D, which 
is above the horizon of the Bowden marl. The geographic range of 
the genus in the West Indies is, therefore, from the Antiguan 
Oligocene to a horizon appreciably above that of the Bowden marl. 

POCILLOPOSA ABNOLDI, new «»e«le0. 

Plate 76, figs. 3, 3a, Zb. 

The type, which is a fragment of a branch, is 28 nmi. long, diameter 
of lower end 6.5 by 12 mm., diameter of upper end 5.5 by 9 mm. 
The cross section of the branch is strongly compressed, and one side 
near and at a place of bifurcation is concave instead of being convex. 
There are no vemicae. 

Calices slightly oblong, lesser diameter about 0.75 mm., longer 
diameter, parallel to the axis of the branch, from 1 to 1.25 mm. 
Cavities rather deep, about 0.5 rom., and steep-walled. Intercoralhte 
areas flattish, arched, or sUghtly crested in profile, of imequal width, 
from 0.3 mm. to 1 mm. across. Coenenchymal surface granulo- 
costulate, graniilations fairly coarse. 

Septa rudimentary, occur as low, blimt-topped, perpendicular 
ridges on the inside of the caUcular walls. In some calices 12 of 
these ridges may be distinguished. The bottom of the cahce is flat 
or very gently concave; no vestige of a colimiella could be found. 

Coenenchyma sohd; coraUite cavities solidly filled except a few 
in the axis of the branch. 

Locality and geologic occurrence. — Canal Zone, station 6444, quarrj^ 
in the Emperador limestone, Empire, collected by Dr. Ralph Arnold, 
whose name I take pleasure in attaching to this weU-marked species. 

Type.— 1^0. 324782, U.S.N.M. 

Of the other five fossil species of PoaUo'pora known from the 
Tertiary formation of the West Indies and Central America, the 
unnamed species from Antigua, previously mentioned, is the most 
similar. The latter species is composed of small, more or less com- 

37149— 19— BuU. 103 11 



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344 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

pressed branches, it has no vemicae, the calices are rather deep, the 
septa are perpendicular ridges down the inside of the calicular walls, 
and there is no trace of a columella. In these characters the two are 
similar. The species from Antigua differs from P. amoldi by having 
larger calices, lesser diameter 1 mm. or more, usually more than 1 mm., 
and the calicular margin is rather persistently marked by a slightly 
raised acute rim. A description of the species from Baracoa, Cuba, 
follows. 

POCILLOPOBA BABACOXBNSIS. new ■ pede j . 

Plate 77, figB. 1, la. 

This species may be characterized as follows: 

The corallum is branching; it has no verrucae and no columellar 
tubercle. The branch is regularly subcircular or broadly eUiptical 
in cross section, 10.5 mm. in diameter at lower end. The calices are 
very shallow and are subcircular in outline, about 0.75 mm. in 
diameter, distance apart usually slightly more than the calicular 
diameter. Thick short septa join the columellar plug to the wall. 
Coenenchyma very dense. 

These characters are different from those of any of the other known 
American species. 

Locality and geologic occurrence. — Cuba, station 3476, in yellow, 
argillaceous marl, Baracoa, associated with Stylophora granulata 
Duncan, collected by T. W. Vaughan. The geologic horizon of this 
species is that of the Bowden marl. 

Type.— No. 324783, U.S.N.M. 

POGILLOPORA GUANTANAMENSIS, mw ■ pede j . 
Plate 77, figs. 2, 2a. 

Corallmn composed of irregularly shaped, more or less compressed 
and contorted branches, among which there is considerable anas- 
tomosis. The branches may be as much as 27 mm. wide, 7.5 mm. 
thick near the summit, and 12 mm. thick at the base. The branch 
on which these measurements were made is 41 mm. long. Verrucae 
entirely absent on the type. 

Cahces from 0.75 to 1.25 mm. in diameter; usually less than or 
about their diameter apart. They are deep pits without any trace 
of septa, except that in a few calices what appear to be thick direc- 
tives are recognizable on the plug forming the calicular floor. CaUcu- 
lar* margins usually even with the coenenchymal surface; in some 
cahces they are somewhat tumid and slightly elevated. 

The columella is only a plug. Stout, horizontal tabulae present. 

Coenenchyma very dense. Surface in type worn, but apparently 
beset with spines or granulations and not costulate. 

Locality and geologic occurrence. — Cuba, station 7514, about 5 
miles nearly duo east of Monument H4 on the east boundary of the 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 346 

U. S. Naval Reservation, Guantanamo, altitude about 400 feet a. t., in 
beds of the age of the Antigua formation, collected by O. E. Meinzer. 
Type.— Ho. 324784. U.S.N.M. This species differs so markedly 
from the other West Indian species of PociUopora that comparisons 
with the other species seem imnecessary. 

Genoa MADRACIS Milne Edwards and Haime. 

1849. Azhelia Milne Edwards and Haike, Comptes Rend., vol. 29, p. 69. 

1849. Madrads Milnb Edwards and ^aimb, Comptes Rmid., vol. 29, p. 70. 

1861. Reussia Duchassaino and Michelotti, M^m. Gorall. Ant., p. 63 (of reprint). 

1871. Pentalophora Savillb-Kent, Proc. Zool. Soc. London for 1871, p. 283. 

1884. Madrads Duncan, Linn. Soc. London Joum., Zool., vol. 18, p. 45. 

1900. Madrads Vauohan, U. S. Geol. Survey Mon. 39, p. 128. 

1901. Axhelia Vauohan, U. S. Fish Commission Bull, for 1900, vol. 2, p. 294. 

1902. Madrads Vbrrill, Conn. Acad. Arts and Sci. Trans., vol. 11, p. 108. 

Type-species. — Madrads asperula Milne Edwards and Haime. 

MADRACIS MIRABIUS (DneluMMliif and MielMlottI). 

1861. Stylophara mirabilis Duchassaino and Michblotti, M6m. Corall. Ant., 

p. 62 (of reprint), pi. 9, figs. 6, 7. 
1901. Azhelia mirahilis Vauohan, tJ. S. Fish Commission Bull, for 1900, vol. 2, 

p. 295, pi. 1, figs. 3, 3a. 

A single fragment of a branch from Limon, Costa Rica, is 23 mm. 
long, 2 mm. in diameter at the lower end, and 3 mm. in diameter 
just below trifurcation at the upper end. The fragment is shghtly 
arcuate in form, not quite straight, and is not so crooked as is usual 
in the specimens of if. mirahUis with which I have compared it. 
The septa are less exsert around the calicular margins then is usual 
in the species. Although there are the differences indicated, they 
are of the kind that may be produced by vegetative causes. 

Locality and geologic occurrence. — Costa Rica, hills of Port Limon, 
No. 669 of H. Pittier collection; geologic horizon not known. 

Cuba, station 3461, gorge of Yumuri River, Matanzas, 19 frag- 
ments collected by T. W. Vaughan in a marl of lower Miocene 
(Bowden) age. 

These fragments perhaps should be referred to a new species; 
but they appear more probably to be only a variant of if. mirahUis. 

Family ASTROCOENIIDAE Koby. 

Genns ASTROCOENIA Milne Edwards and Haime. 

1848. Astrocoenia Milne Edwards and Haime, Comptes Rend., vol. 27, p. 469. 
1900. Astrocoenia Gregory, Palaeontol. Indica, eer. 9, vol. 2, pt. 2/ p. 59. (Syn- 
onymy and elaborate discuaeion.) 

Type-species. — Astrea numisma Defrance. 

Besides the iive species of Astrocoenia recognized in the present 
paper, I have described one under the name of Stylocoenia duerdeni 



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346 BULLETIN 103, UNITED STATES NAZIOKAL MUSEUM. 

from the Ek>cene of Jamaica,^ which also occurs in the upper Elooeae 
of St. Bartholomew. I describe as new the species from Antigua 
{A. deccUurensis), to which Duncan applied the name Astroeoerda 
amata.^ This species is also found in the coral reef at the base of the 
Chattahoochee formation on Flint River, near Bainbridge, Georgia, 
and near Guantanamo, Cuba. More critical -study may lead to the 
recognition of one or two additional species. The names of all 
European species applied by Duncan and others to West Indian 
forms probably should be dropped from the literature. 

ASTBOCOENIA D'ACHUBDn Duwu. 

Plate 78, figs. 2, 2a. 

1873. Astrocoenia (TaehiardH Duncan, GeoL Soc. London, Quart. Joum., voL 29, 

p. 554, pL 20, figs. 7, 7a. 
1899. Astrocoenia cTachiardii Vauohan, Mus. Comp. Zool. Bull., vol. 34, p. 229. 

Dr. C. W. Hayes obtained in Nicaragua, "on or near the Pacific 
coast," a specimen of Astrocoenia (pi. 78, figs. 2, 2a) that seems 
referable to A. d*achiardii. 

The corallum is ramose; branch somewhat compressed, lesser 
diameter of lower end 10.5 mm., greater diameter only slightly more 
than the lesser. 

Calices from 2 to 3 mm. in diameter, measured between thecal 
summits; the diameter of the largest calice is 3 mm. Maximum 
thickness of walls between adjoining calicular cavities, 1 mm. 
Depth of caUces about 1 mm. 

Eight prominent septa reach the columella, with a small septum 

between each pair of the larger. The large septa are narrow above 

the bottom of the calice, where they widen and fuse to the columella, 

aroilnd which they show decided thickening. The calicular cavity, 

. therefore, is steep-sided and relatively flat-bottomed. 

The columella is a slightly prominent, compressed style. 

Locality and geologic occurrence. — Nicaragua, on or near the Pacific 
coast, in the Brito formation, collected by C. W. Hayes. Dr. Hayes 
says regarding the Brito formation.' 

The greater part of the Brito formation is apparently banen of oiganic remans. 
The only location at which foaails have been found are on or near the Pacific coast. 
This, however, may be due to the fact that the rock exposures are not elsewhere of 
such a character as to facilitate the discovery of fossils, and the latter may possibly 
be more generally distributed than present knowledge would indicate. The fossils 
are confined almost wholly to the limestones and marly beds. They consist of coials, 
molluscan, and foraminiferal remains. 

The Foraminifera, according to Dr. Joseph A, Cushman, indicate 
an Eocene age. 

1 Mus. Comp. Zool. Bull., voL 34, p. 235, pi. 87, figs. 1-4, 1899. 

* Qeol. Soc. London Quart. Journ., toL 19, p. 425, pi. 14, fig. 7, 1863; Idem., vol. 2t« p. SS, 1867. 

* GeoL Soc. Amer. Bull., voL 10, p. 312, 1899. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 347 

Astrocoenia d^acMd/rdii was described from the upper Eocene of St. 
Bartholomew. Finding it on the Pacific coast of Nicaragua is addi- 
tional evidence in favor of connection between the Atlantic and Pacific 
oceans across Central America during upper Eocene time. 

ASTBOOOENU GUANTANAMBNSIS, new i tede fc 

Pkte 79, figs. 1, la, 2. 

CoraUum massive, with a rather imiformly rounded or more or less 
tuberose surface. Type 55 mm. long, maximimi width about 31 mm., 
height 38 mm. The corallum may be much larger. 

Calices polygonal, shallow, almost superficial, small; maximum 
size about 1.75 mm. in diameter, 1.5 mm. usual, smallest calices 
about 1 mm. in diameter, measured between thecal summits. Inter- 
corallite walls acute or flattish, usually less than 0.25 mm. wide, 
maximum width 0.5 mm. ; crossed by subequal costae corresponding 
to aU septa imless very narrow, when the edge of the waU is dentate 
instead of costate. 

Septa 16 in number, 8 reach the columella; 8 small, about half 
the length of the principals; in most instances they are thicker in 
the wall than at their inner ends. Margins of the longer with about 
three dentations on each. Septal faces with sharp granulations. 

Columella, a small, erect, central style. 

Localities and geologic occurrence. — Cuba, station 7522, Mogote 
Peak, 0.5 mile east of east boundary of United States Naval Reser- 
vation, Guantanamo, south side of peak, altitude about 375 feet a. t., 
collected by O. E. Meinzer (type). 

Antigua, station 6865, Jackass Point, St. John, collected by T. W. 
Vaughan. 

Panama, station 6587, Tonosi, collected by D. F. MacDonald. 

ryp6.— No. 324794, U.S.N.M. 

Astrocoenia guarUanamensis is most nearly related to Astrocoenia 
incrustans (Dxmcan) which is from the upper Eocene St. Bartholo- 
mew limestone, and is the next species here described. The calices 
of A. incruatanSf a description of which follows, are rather deep and 
the intercorallite areas are flattish and costate. 

ASTROCOENU INCBUSTANS (Dimam). 

1873. Stephanocoenia inerusUms Duncan, Geol. Soc. London Quart. Journ., 

vol. 29, p. 653, pi. 20, fig. 6. 
1899. Stephanocoenia inenutane Vaughan, Mus. Comp. Zool. Bull., vol. 34, 

p. 229. 

Original description. — '*The corallum is low in height, and incrusts 
rocky surfaces. The coraUites are united by their rather thick walls, 
and are parallel. The calices are quadrangular or pentangular, and 
their margins are marked by the septa of the adjacent coraUites. 



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348 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

The septa are subequal at the wall, and 16 in number; but only 
eight reach the small and deep styloid columella; the others project 
very slightly, and are moniliform on their free edge. The paU are 
attached to the eight larger septa. 

"Height of corallum, one-tenth inch [ = 2.5 mm.]. Breadth of 
calice, one-twentieth inch [ = 1.25 mm.]." 

The following notes are based on the type-specimen: 

It is a small thin fragment, 17.5 mm. long, 8 mm. wide, and 4 mm, 
thick. 

The calices are moderately deep polygonal, many are elongate, the 
smaller ones measure 0.9. mm. in diameter, an elongated one is 1.2 
mm. wide and 2 mm. long. The walls are thin, about 0.2 mm. wide; 
however, the upper edges of the septa are flattened and somewhat 
expanded. No mural styles. 

Septa, 16 in number, equal in thickness at the wall, thicker than 
the spaces between; 8 extend to the coliunella, the laminae thinner 
between the portions surrounding the columella and the outer ends. 
The other 8 septa are short. The margins are finely dentate. Dis- 
tinct pali absent. Apparently dissepiments are present. 

Columella styliform, rather prominent, compressed. 

This coral can not be referred to Steph-anocoenia because there are 
no pali and the septal margins are dentate, instead of being entire. 
However, it exhibits all the characteristics of Astrocoenia. In the 
size of the calices, number of the septa, and character of the septal 
margins it resembles A, duerdeni (Vaughan), but differs from that 
species by the apparent absence of mural spines Notwithstandmg 
this, it is not impossible that the type-specimen could be a portion 
of a corallum of A, duerdeni ^ the styles being absent from the area 
whence it was derived. 

Locality and geologic occurrence. — ^Island of St. Bartholomew, 
P. T. Cleve, collector; subsequently collected by T. W. Vaughan; in 
the upper Eocene St. Bartholomew limestone. 

Type, — ^University of Upsala. 

ASTSOCOBNU DBCATURBNSIS. Mw ted e iu 
Plate 78, figs. 3, 3a, 4, 4a. 

1863. Astrocoenia ornaUi Duncan, Geol. Soc. London Quart. Jonm., vol. 19, 

p. 425, pi. 14, fig. 7. (Not Milne Edwards and Haime.) 

1864. Astrocoenia omaia Duncan, Geol. Soc. London Quart. Joum., vol. 24, p. 23. 

Corallum massive, rather lai^e, upper surface with numerous gib- 
bosities. One specimen has a base 14 by 17 cm., respectively, as the 
smaller and greater diameter, and is about 8 cm. in height, another 
has 19 cm. as the greatest diameter of the base. 

CoraUites polygonal, separated by walls that are never very thick, 
rarely as much as 1 mm., upper edge usually if not always marked 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 349 

by a small raised, granulated line. The distal ends of the septa are 
produced as short costae to this line and often a granulation occurs 
between each pair of costae. The diameter of the coraUites ranges 
from 1.5 to 2.5 mm. ; about 2 nam. is the average. Calices shallow. 

Septa distant, normally 16 in number, of which 8 extend to the 
columella; occasionally 20, with 10 reaching the columella. Their 
outer ends are slightly prominent on the wall and are equal in size. 
The inner margins lie almost in a straight line or are very slightly 
excavated but are regularly finely dentate, with four to seven teeth 
to each septum. These teeth are moderately acute and are directly 
obliquely upward and inward. Granulations on the faces minute, 
pointed. 

Endothecal dissepiments present, thin, not abundant. 

Columella a strong style, upper end pointed but not very promi- 
nent. There is some thickening of the inner ends of the larger septa 
where they fuse to the columella. 

Localities and geologic occurrence. — Geoi^a, station 3383, Hale's 
Landing on Flint River, 7 miles below Bainbridge; and station 3381, 
Blue Springs, 4 miles below Bainbridge, collected by T. W. Vaughan. 

Island of Antigua, West Indies, in the Antigua formation, collected 
by Robert T. Hill. 

Cuba, station 7523, south side of Mogote Peak, altitude 250 feet 
a. t., one-half mile east of east boundary of the United States Naval 
Reservation, near Guantanamo, collected by O. E. Meinzer. 

Type.— CAt. No. 324789, U.S.N .M. 

Paraiype.—CeA. No. 324788, U.S.N.M. 

Astrocoenia omaia Dxmcan from Antigua (No. 12948, coll. Geol. 
Soc. London) is a massive species of Astrocoenia. It is silicified; the 
coraUites are crowded, polygonal, intervening walls thin, diameter of 
corallites, 1.5 to 1.75 mm. Septa, 8 principal; 8 rudimentary, thin 
and distant. Columella, a slender style. 

ASTROCOENIA MEINZERI. new species. 
Plate 7&, figs. 3, 3^. 

Corallum composed of thick branches, with broadly eUiptical cross- 
section. Type, a broken, bifurcating branch. Length from broken 
base to fork, 50 nmi.; diameter of basalend, 23.5 by about 24 nmi. 
Diameter of broken end of branch at fork, 23 by 24 mm. Length of 
broken lateral branch from fork, 21 mm.; diameter of distal broken 
end, 17.5 by 20.5 mm. 

Calices rather large, diameter measured between thecal simimits 
from 2.5 to 3 nmi.; depth, 1.25 to 1.5 mm. Intercorallite walk from 
0.5 to 1.5 mm. across where well preserved, about 0.75 mm. usual. 
In places the top of the wall is acute, but this condition is probably due 



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350 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

to weathering. Where the walls are wide there is usually a distinct 
intercorallite groove. Thick costae or mural teeth are probably 
present on perfect spedmens, but they are not distinct on the type, 
as its surface is worn. 

Septa Id in number; 8 principals extend to the columella, and 8 
are short but thick. The {urincipal septiEt slope in a concave curve to the 
bottom (A the calioe, and are narrow nearly to the level ci thb bottom 
of the calice; the smaller septa are narrow. All septa are thick in 
the waU, and the principals are fused by their thickened inner ends 
around the columella. About seven small dentations were counted 
on one long septum. Septal faces with small granulations. 

Columella a low style, with rounded upper end; it with the inner 
septal ends fused around it forms a rather large columellar mass. 

Thickish dissepiments are present. 

Locality and geologic occurrence. — Cuba, station 7522, Mogote Peak, 
0.5 mile east of east boundary of United States Naval Reservation, 
Guantanamo, south side of peak, altitude about 375 feet a. t., col- 
lected by O. E. Meinzer. 

Type.— No. 324791, U.S.N.M. 

The species most nearly related to Astrocoenia meinzeri is A. 
d^achiardii Dimcan from the upper Eocene St. Bartholomew lime- 
stone. The branches of A. d^dckiardU are more irregidar in form, 
for the same size branch the calices are larger, up to 3.6 mm. in 
diameter, the intercorallite walls are not so wide, the outer part of 
the septal margins are steeper, and the septal dentations are coarser. 
Notwithstanding these apparent differences, it should be admitted 
that larger collections may lead to combining the two supposed 
species. 

ASTBOCOBNIA POBTORICENSIS, new Bpedes. 
Plate 76, figs. 4, 4a; plate 78, figs. 1, la. 

1901 . Astrocoenia omata Vaughan, Geol. Soc. London, Quart. Jour., vol. 57,p. 497 " 
Not: 
1838. Pontes omata MiCHELom, Specim. Zooph. diluv., p. 172, pi. 6, Gg. 3. 
1857. Astrocoenia omata Milne Edwards and Haime, Hist. nat. Corall., vol. 2,p. 
257 
The following is a description of the type (pi. 76, figs. 4, 4a) : 
Corallum forming flattened, even palmate branches. The type- 
specimen, which is broken, has a greatest width of 53 mm., length 
105 mm., and a thickness of 15.5 mm. at the lower and of 7.5 nmi. at 
the upper end. 

Calices, diameter from 1.0 to 1.5 mm., excavated but rather shallow, 
outline polygonal, united by compact, rather narrow walls, which 
range from 0.2 to 0.5 mm. across. The distal ends of the septa form 
low costae. 

Septa, 16 in number, 8 reach the calumella and 8 are short or even 
rudimentary; a few dentations, usually about 3 or 4 on the margin 



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GEOLOGY AND PALEONTOLOGY OF THE OANAL ZONE. 351 

of each principal septum. Interseptal loculi about as A\ide as the 
thickness of the septa. 

OoliuneUa an erect style, which does not reach the level of the upper 
edge of the wall; its upper termination rounded; cross-section eUip- 
tical. 

!Endothecal dissepiments present. 

JjOcdlitieB and geologic occurrence. — Island Antigua in the Antigua 
formation. Collected by R. T. Hill and by T. W. Vaughan. 

Porto Rico, station 3191, 4 miles west of Lares, in the Pepino forma- 
tion, collected by R. T. Hill. 

Canal Zone, station d024&, in the Emperador limestone, at the 
crossing of the Panama Railway over Rio Agua Salud, collected by 
T- W. Vaughan and D. F. MacDonald. 

Type,— No. 324785 U.S.N.M., from 4 miles west of Lares, Porto, 
Pepino formation, collected by R. T. Hill. 

Paraiype.— Cat. No. 324786, U.S.N.M. 

The foregoing description is based on the type-specimen and does 
not take into consideration the variation of the species. I obtained 

6 good suite of specimens at two exposures of the Antigua formation 
on the island of Antigua. The branches range in form from greatly 
compressed to subcylindrical (see pi. 77, figs. 1, la, illustrations of a 
specimen from Willoughby Bay, Antigua). A segment from near 
the base of a subcylindrical branch was collected on Rio Agua Salud, 
Canal Zone. 

Genus STTLOCOENIA Milne Edwards and Haime. 

1849. Stylocoenia Milsb Edwabds and Haoie, Gomptes Rend., vol. 27, p. 469. 
Type-species. — Asirea emarciata Lamarck. 

STTLOGOENU PUMPELLYI (Vangluui). 

1900. Astrocoenia pumpellyi Vaughan, V, S. Geol. Survey Mon. 39, p. 149, pi. 
17, figs. 7, 7a. 

This species seems to belong to the genus Stylocoenia, as it has inter- 
corallite pillars; but as some septa show dentations on their margins, 
the original generic identification may be correct. It occurs in the 
base of the Chattahoochee formation, near Bainbridge, Georgia, and 
not in ^cksburgian deposits, as I stated in the original description. 

Localities and geologic occurrence. — Georgia: Station 2326, Russell 
Spring, Flint River, 4 miles below Bainbridge, collected by R. Pum- 
pelly (type. Cat. No. 158315, U.S.N.M.); station 3381, same locality 
as the preceding, collected by T. W. Vaughan; stations 3383, col- 
lected by T. W. Vaughan, and 7078, collected by T. W. Vaughan, 
C. W. Cooke, and W. C. Mansfield, Hales Landing, Fhnt River, 

7 miles below Bainbridge, in the base of the Chattahooche formation. 



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352 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Antigua: Station 6881, Willoughby Bay, collected by T. W. 
Vaughan in the Antigua formation. 

Family OCULINIDAE Milne Edwards and Haime. 

Genus OCULINA Lamarck.* 

1816. Oculina Lamarck, Hiflt. nat. Anim. sans Vert., vol. 2. p. 283. 

1849. Oculina Milne Edwards and Haime, Comptes Rend., vol. 29. p. 68. 

1850. Oculina Milnb Edwards and Haime, Men. Brit. foes. Cor.. Intr., p. XIX. 

Type-species. — Madrepora virginea Ellis and Solander. 

OCUUNA DIFFUSA LMBarck. 

1816. Oculina diffusa Lamarck, Hist. nat. Anim. sans Vert., vol. 2, p. 285. 

1901. Oculina diffusa ? variety Vauohak, U. S. Fish GommisBion BuU. for 19U0, 

vol. 2 p. 294, pi. 1, figs. 5, 5a. 
1915. Oculina diffusa Vauohan, Washington Acad. Sci. Joum., vol. 5, p. 596. 
1915. Oculina diffusa Vaughan, Carnegie Inst. Wash. Yearbook No. 14. p. 227. 

Doctor MacDonald obtained seven pieces of branches of this 
species at the locahty mentioned below. They are slender and 
resemble fragments from specimens of OcuXina diffusa, which grow 
either in water 10 to 16 fathoms deep or where the water is very 
quiet. The specimens from Panama nearly duplicate those I de- 
scribed from Porto Rico. 

Locality and geologic occurrence, — Canal Zone, station 5849, swamp, 
Mount Hope, Pleistocene, collected by D. F. MacDonald. 

OCUUNA VARICOSA LeSneiir. 

1820. Oculina varicosa Le Sueur, Paris Mus. M^., vol. 6, p. 291, pi. 17, fig. 19. 

1902. Oculina varicosa Vbrrill, Conn* Acad. Sd. Trans., vol. 11, pi. 82, figs. 2. 3, 

4 (refs. to literature). 

A single nearly typical fragment of a branch was obtained. 
Locality and geologic occurrence. — Canal Zone, station 5849, swamp, 
Mount Hope, Pleistocene, collected by D. F. MacDonald. 

ARCHOHELIA, new genus. 

ArcTiohelia differs from Ocuiina solely by having a persistent axial 
coralUte, whereas in OcvMna there is no axial corallite. Pali or pali- 
f orm teeth are present on all but the last cycle of septa. Columella 
trabecular, with some papillae on its upper surface. 

Type-species. — Archyhelia limonesis Vaughan. 

The relations of this genus to the species described in my mono- 
graph on the Eocene and lower Oligocene coral f aimas of the United 
States ^ under the names Astrolielia neglectaj A. bumsi, Oculina vicJcs- 
burgensiSj 0. mississippiensis, 0. singUyi, 0. aldbamensis, 0. harrisi. 

1 Toula (K. K. Geolog. Relohsaiistalt Jshrb., vol. 61, p. 489, pi. 30, fig. 1, 1011) applies the name Ocm- 
Utta gatunentii to a pieoe of a branch of coral, but his descrlptioxi and figure are inadequate for the ides- 
tiflcation of the species. 

t U. S. Oeol. Survey Monograph 89, pp. 114-124, 1900. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 



353 



O.aMrichi, and 0. ? smithi should be indicated. The species mentioned 
have axial corallites and genericaily resemble ArchoheUa except in 
the details of the inner ends of the septa. The type-species of 
AstrTislia (the correct spelling of the name, instead of Aatrohelia) is 
Madrepora paimaia Goldfuss; which has no definite axial corallites^ 
and I have seen no pah or paliform lobes on its septa. The species 
to which I appUed the names Astrohdia neglecta and A. bumsif as 
they possess axial coralhte should be taken out of the genus 
AstrMia. As it is not practicable just now to revise critically the 
Ek>cene and lower Oligocene species hsted above, it will here only be 
mentioned that they probably should be transferred to Archdhdia. 

ARCHOHEUA UMONENSIS, new spedee. 

Plate 80, figs. 1, la, 16, 2, 3. 

Coralliun composed of relatively slender branches. The following 
are measurements of the cotjrpes: 

DimsrmoTU in millimeters ofcotypes of ATchohelia limontrms. 





Branch. 


Length. 


Diameter. 


Calioes 






Lower 
end. 


Upper 
end. 


1 
Diameter, i 


Exsort. 


1 


25 
31 
33 


4.5 
4.5 
4 


4 

4 
3.5 


2.6-3 1 

2-2.6 

2.3-3 


1-3.5 


2 


0.5-3.6 


3 


1-7 







The cavity of the axial corallite is about 2.25 in diameter. The 
forgoing tables give the dimensions and amount of the projection, 
of the radial calices — the diameters stated are as measured from the 
outside of the walls. The distance between adjacent calicular mar- 
gins is about 2.5 mm. on branch No. 2; in extreme cases it ranges 
up to as much as 7 mm., as between some calices on branch. No. 3. 
The arrangement is in more or less definite spirals. Subequal or 
slightly alternating costae, with closely granulate surfaces, cor- 
respond to all septa just below the caUcular edges; lower down on 
the corallite limbs they flatten and become subequal; they may 
continue on the coeneuchymal surface or disappear. The cahcular 
cavities are excavated; moderately deep, about 1.5 mm. 

Septa normally in three complete cycles; primaries as a rule slightly 
larger than the secondaries, both cycles reach the columella, and 
have subequal, slightly exsert upper margins; tertiaries smaller than 
the secondaries and have lower upper margins. Inner edges of the 
tertiaries usually free, but in some systems they fuse to the sides of 
included secondary septa. Single or double paliform teeth on the 
inner ends of the primaries and secondaries. Septal faces closely 
granulate. 



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354 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Columella papillate. 

Coenenchyma dense; with or without costal prolongations from 
the calicular peripheries; fine granulations scattered over its surface. 

LocdliHes and geologic occurrence. — Costa Rica, limon, as fdlows 
Station 2692, coUected by R. T. HiU; Moin HiU, Niyeau d and No. 
461, collected by H. Pittier; station 58846, Moin Hill, coUected by 
D. F. MacDonald. The geologic horizon seems to be PlioceiiA. 

Florida, station 3300 in the Pliocene Caloosahatchee marl of Shell 
Creek, coUected by Frank Bums. 

Ootypes.—Yio. 324809, U.S.N.M., from Niveau rf, Moin HiU, Port 
Limon (3 specimens). 

Family EUSMILIIDAE VerriU. 

Genus ASTEROSMILIA DaBcan. 

1867. AUerotmUia Duncan, Roy. Soc. Philos. Trans., vol. 167, p. 653. 
1873. Aiierosmilia Duncan, Geol. Soc. London Quart. Joum., vol. 29, p. 553. 
1884. AsterosmUia Duncan, Linn. Soc. London Joum. Zool., vol. 28, p. 61., 

Type-Species. — TrochocyatJivs abnormdlia Duncan. 

When Duncan described this genus ho referred to it his Trocho- 
cyathua aJmormdlie, changing the name to anomdUif and refigured 
the species. He also described two additional species as Astero- 
smUia exarata and A. comtUa, a synonym of A. abnormaUs, and 
faUed to designate a type-species for the genus. Trochocyathus 
cbbnormoLia was described with much care, while the descriptions of 
the two other species are short and unsatisfactory. A. comuta is a 
synonym of A. dbnormalis. It therefore seems best to take the 
species I have selected, as indicated above, as the type-species of 
the genus. 

Duncan described three species of Asterosmilia from the Tertiary 
formations of Santo Domingo, namely, Troc'hocyai'huB ahnormalis^^ 
for which the genus Asterosmilia was subsequently erected, A. car-- 
rmta, and A. exarata j^ and one species A, pourtalesi from the upper 
Eocene St. Bartholomew Umestone. I consider A, comuta a syno- 
nym of A. ahnormdlis, and transfer Duncan's Trochocyaihus profundus 
from the genus in which it was originally placed to AsterosmUiay 
leaving four described fossil species in the genus. Pourtalfes described 
from the West Indies one recent species that belongs to Asterosmxliaj 
his A, prolifera, originally named Ceratocyathus prclifer, and of which 
Lindstrom's Para^cyathus arcuaius is a synonym. I here describe an 
additional new species, namely, A. JiiUi, from Bowden, Jamaica, and 
Ldmon, Costa Kica, and have described two additional species from 
Santo Domingo, in a paper not yet published, making eight, the total 
number of American species at present known to belong to the genua 

I Collected by A. Olsson on Provision Island, Costa Rica, in the Qatun formation. Footnote added 
to page proof. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 



355 



ASTBBOSMIUA HILU, new 

Plate 80, figs. 4, 5, 6, 6a. 
1899. Asterotmilia species Yaughan, Mus. Gomp. ZooL BulL, voL 34, p. 149. 

I find it difficult to explain why a species so common as this one 
cotild have so long remained midescribed. There are from Bowden, 
Jamaica; 41 specimens in the Henderson and Simpson collection, 20 in 
the Hin collection, and 9 in the T. H. Aldrich collection, making 
a total of 70 specimens that I have studied from this one locality. 
A series of ten of the best specimens of the Henderson and Simpson 
collection have been selected as the cotypes. 

Corallum comute with a pointed base and attached, at least in its 
early stages, rather slender, curved in the plane of the greater trans- 
verse axis of the calice. The following table gives the measurements 
and ntunber of septa in the type specimens. 

Dimensions of and number of sepia in Asterosmilia hilli. 



8])ecimexi No. 



1- 
2. 
3. 

4. 
5. 
6. 

7. 

S. 

9. 

10. 



Greater 
diameter 


diameter 


Heisht of 
corsulum. 


of calico. 


ofcalioe. 


mm. 


mm. 


mm. 




3 


6.5 




3.75 


7 




3.6 


9 


4.5 


4 


10.5 


e.6 


5.5 


12 




6 


15 




5 


15.5 




7 


18.5 


0.6 


8.75 


19 


>10 


19 


25 



Number of septa. 



About 24, and probably some rudimentary. 

24, and a few rudimentary. 

24, and a few rudimentary. 

24+17 of the fourth cycle. 

24 +20 of the fourth cycle. 

244-20 of the fourth cycle. 

Calloe broken on side. 

24+20 of the fourth cycle. 

48, fourth cycle complete. 

48, four complete cycles. 



1 About. 



The caUce is obhque, its upper edge being considerably higher than 
its lower. In the measurements given above the height of the coral- 
limi is measured from the tip of the pedicel to the highest point of the 
calicular margin. 

The wall is only moderately thick, externally there is a variable 
amoimt of pellicular coating. Costae corresponding to all septa, 
distinct, but usually not prominent. There is a fair amount of varia- 
tion in the costal characters. In some specimens the costae of all 
cycles are equal or subequal, low, flattish or only slightly crested; 
in others, those corresponding to the septa of the first and second 
cycles of septa are decidedly more prominent than the intervening 
costae. Those corresponding to the third cycle" of septa may be 
slightly more prominent than those corresponding to the fourth. 
Sometimes costae of both kinds are combined in one specimen. 
Rather often in an intercostal space there is a raised thread or line 
which does not correspond to a septum. Minute, crowded granula- 
tions are scattered over the surfaces of the costae and in the inter- 
costal spaces. 



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356 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Septa; thin, distant, those of the first and second cycles have slightly 
exsert margins. In adult specimens, 19 to 25 mm. tall, there are 
four complete cycles, in younger specimens the fourth cycle is in- 
complete. The members of the first and second cycles are of equal 
size, extend to the columella, and are decidedly thicker than the other 
septa. The members of the fourth cycle are thinner and shorter than 
those of the third. The septal margins are subentire, arched above 
and fall at a very steep angle to the bottom of the calicular fossa. 
Septal faces finely striate, with more or less elongate granulations 
along the courses of the striae. Line of divergence of the striae very 
dose to the inner side of the wall. Wide, tall, thin, pali, rounded 
above, stand before the septa of the third cycle, from whose inner 
margin they are separated by a deep notch. The width of a palus is 
about 1 mm., height, 1.5 mm. 

Dissepimental endotheca, present, but not abundant. The dis- 
sepiments thin. 

The columella in fully grown specimens, prominent, compressed or 
even distinctly lamellar in appearance. In young and broken speci- 
mens it appears to be composed of interfused process^ from the 
inner ends of the septa, it is decidedly vesicular. CaUce, radier 
deep, 3 to 4 mm. 

Localities and geologic occurrence, — Jamaica, Bowden, collected by 
J. B. Henderson and C. T. Simpson and R. T. Hill. 

Costa Rica, ''CoUine en demolition," Limon, Costa Rica, No. 618 
of H. Pittier's collection. 

Cotypes,—i^os, 324815, 324816, U.S.N.M. (10 specimens). 

The specimens from Limon, Costa Rica, are essentially duplicates 
of those from Bowden. One specimen with a greater calicular diam- 
eter of 9.5 mm. has a few quinary septa. 

Genus STEPHANOCOENIA Milne Edwards and Haime. 

1848. Suphanocoenia Milne Edwards and Uaime, Gomptee Rend., vol. 27, p. 469. 
1848. Suphanocoenia Milne Edwards and Haim e, Ann. Sci. nat., Zool., ser. 3, 

vol. 10, p. 300. 
1850. Stephanocoenla Milne Edwards and Haime, Mon. Brit. foes. Cor., Intr., 

p. XXX. 
1857. Suphanocoenia Milne Edwards and Haime, Hist. nat. Gorall., vol. 2, 

p. 264. 
1884. Antillastraea Duncan, Linn. Soc. London Joum., Zool., vol. 28, p. 108. 

Type-species, — Astrea intersepta Lamarck = Madrepora intersepta 
Esper. 

STEPHANOCOENU INTERSEPTA (Esper). 

1795. Madrepora intersepta Esper, Pflanzenth., Fortsetz., p. 99, pi. 79, figs. 1-3. 
1816. Astrea intersepta Lamarck, Hist. nat. Anim. sans Vert., vol. 2, p. 266. 
1848. Suphanocoenia intersepta Milne Edwards and Haime, Comptes Rend., 
vol. 27, p. 469. 



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GEOLOGY Ai^D PALEONTOLOGY OF THE CANAL ZONE. 357 

1848. Suphanocoenia inursepla Milne Edwards and Haime, Ann. Sci. nat., 

ser. 3, ZooL, voL 10, p. 300, pi. 7, figs. 1, lo, 16. 
1848. Suphanocoenia michelinii Milne Edwards and Haime, Ann. Sci. nat., 

ser. 3, ZooL, vol. 10, p. 30L 
1864. Plesiastraea spongiformis Duncan, GeoL Soc. London Quart. Joum., vol. 20, 

p. 39, pL 4, figs. 6a, 66. 
1866: Suphanocoenia debilis Duchassaing and Mighblotti, Sup. M6m. CoralL 

Antilles, p. 76, pL 9, figs. 7, 8. 
1884. AntUlastraea spongiformis Duncan, Linn. Soc. London, Joum., ZooL, 

voL 18, p. 108. 
1895. Suphanocoenia inUrsepta Gregory, Geo!. Soc. London Quart. Joum., vol. 

51, p. 276. 
1900. iStep^nocoentatn£«rMptaVAUGHAN, U. S.Geol. Surv. Mon. 39, pp. 152,153. 

1900. Pletiastraea goodei Vbrrill, Conn. Acad. Arts and Sci. Trans., vol. 10, 

p. 553, pi. 67, ^g. 1. 

1901. Suphanocoenia intersepta Vaughan, Geol. Reichs. Mus. Leiden Samml., 

ser. 2, vol. 2, p. 20. 

1902. Flesiagtraea goodei Verrill, Conn. Acad. Arts and Sci. Trans., vol. 11, 

p. 106, fig. 1, p. 172, pi. 31 (not pi. 30 as given in the text), figs. 1, la. 
1916. Suphanocoenia inuraepta Vaughan, Carnegie Inst. Wash. Yearbook No. 13, 

p. 222. 
1916. Suphanocoenia inUrsepta Vaughan, Carnegie Inst. Wash Yearbook No. 14, 

p. 221. 

Although the original description of Lamarck is brief, it is good. 
According to him, "Cette espdce forme de large plaques im peu 
convexe, et oflfre k sa surface un r^eau assez fin, constitu6 par les 
bords r^unis des cellules. On voit im petit axe au centre de chaque 
6toile." He placed Madrepora intersepta Esper doubtfully in its 
synonymy. Esper says regarding his specimens of the species: 
"Els kommt diese Koralle von den ostindischen Meeren; ich habe sie 
gleichfalls durch die Gtite des Herm Prediger Chemnitz, mitgetheilt 
erhalten.^' It appears that Chemnitz had specimens from both the 
Atlantic and the Indo-Pacific and that he gave numbers of them to 
Esi>er. Apparently in some instances the locaUty labels were con- 
fused, and that this is one of them, for Esper^s figures (pi. 79, figs. 
1-3) are fairly good for the West Indian and Floridian species to 
which the specific name intersepta is now apphed, and seem to me 
to represent no other Uving species of coral with which I am f amiUar. 

The corallum is massive, either subhemispherical or pulvinate in 
form. The coralUtes are not protuberant, joined directly by their 
walls or by coetae, in the latter case exothecal dissepiments may be 
present. The diameter of the caUces ranges between 2 and 3 mm. 
Septa in three cycles. Primaries and secondaries bear well-developed 
pali; by which liiey are joinpd to the columella. Tertiaries thin and 
relatively short. Septal margins subentire or very finely dentate. 
Columella, a compressed style of nearly the same height as the pali. 
Endothecal dissepiments subhorizontal, thin, average about 0.5 mm, 
apart. 



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3&8 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

As this is the type-speciee of the genus StepJumocoemay the following 
notes on its finer structure will be repeated, with slight emendation, 
from my paper on the Eocene and lower Oligocene corals of tiie 
United States (1900) : The septa are composed of ascending trabeco* 
lae; near the wall is a line of divergence. External to this line die 
trabeculae pass upward and have a slight inclination outward. The 
trabeculae on the inner side of the line of divergence pass upward and 
incline inward. The trabeculae are fine, measuring from 0.027 to 
0.04 mm. across. A study of the lines of growth across the trabeculae 
indicate an entire or very obscurely dentate septal margin. The 
growth segments of the septa are weU defined; the distance across 
one measured along the line of divergence is about 0.32 mm. on an 
average. The distal ends of the septa do not thicken sufficiently to 
form a pseudotheca. In places dark centers or a dark band can be 
seen in the theca between the septal ends; that is, the wall belongs 
in the euthecal class. In some instances the wall is clearly formed 
by peripherally placed dissepimenta. The corallites are ratlier ottm 
joined by their costae. In such instances the wall of one corallite is 
usually formed by dissepiments. There is usually distinguishable a 
central erect piece, aroimd which the principal septa fusa by their 
inner margins. In some instances the columella appears to be formed 
merely by the fusion of the septal margins. In one caUce the axis 
of the coliunella is vacant, the septal margins having fused around it. 
The pali in cross section show as thickenings on the inner septal ends. 
The inner ends of the tertiary septa are free. 

The above description should be compared with Felix's description 
of Stephanocoenia formosa (Goldfuss).* I should also like to call 
attention to a statement by Miss Ogilvie, that "it is doubtful if tiiey 
{Astrocoenia and StepMnoeoenia) are represented in recent seas."* 
She evidently did not know that the type-species of Stephanocoema 
is the recent 8, intersepta (Esper). So if there is any doubt, it is 
that the genus is found fossil earlier than late Tertiary. 

It is astonishing to find the following statement in a recent pap^ 
by Felix:' "Von dieser Art, welche heutzutage in Australischen 
Meeren lebt, liegen mir zwei examplare vor. Fossil findet sich in dem 
Pliocanen Mergel von Rangoen auf Java." Such a statement when 
the species he is discussing is one of the most widespread and besi 
known of those in Pleistocene deposits adjacent to and in the Recent 
waters of the western Atlantic Ocean, the Caribbean Sea, and the 
Gulf of Mexico ! 

Synonymy. — Gregory in 1895 gave full references to the literature 
on this species up to that date, except that he did not place Stephmo- 
coenia dehilis Duchassaing and Michelotti in its synonym. 

^ Deutsch. Qeolog. Gesell. Zeitschr., vol. 50, pp. 252-26i, pi. 2, fig. I. 

s Roy. Soc. London Trans., vol. 187, p. 307, 1896. 

* KOnlgl. S&cbs OeseU. Wiss., Leipzig, Math. Phys. Kl., vol. 64, p. 444, 1012. 



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GEOLOGY AND PALEONTOLOGY OP THE OANAL ZONE. 359 

While in Turin in 1897 I examined the specimens identified by 
Duchassaing and Michelotti as Stephanoeoenia irUersepta and S. miche- 
Km. They belong to the same species. It is said of jS. ^e&iZt«; "Bien 
que les dimensions des cahces de cette espdce soient les mftmes que 
dans la Stephanoeoenia michelini, elle s'en distingue pourtant par la 
muraille, par les cloisons plus minces, et par les palis qui atteignent 
la hauteur de la colimielle." The only character of apparent value is 
the height of the pali, which are as tall as the columella. The pah 
and columella are usually of nearly the same height in the species; 
in areas on some specimens the colimiella is somewhat taller; in other 
areas the pah are taller. 

I examined Duncan's type of Plesiastraea [later described as AntUr 
lastraed] spongiformis and a specimen identified by him ab Stephano- 
eoenia intersepta. The coraUites of the former are united by their 
costae, and where the costae meet there is often a second Wall outside 
the true coraUite wall. The second specimen had been out, the larger 
piece bearing the label Stephanoeoenia intersepta; the smaller piece, 
which fits into the larger, was labeled Plesiastraea spongiformis: 
Duncan, it seems, could not distinguish between the two. I agree 
with Gregory in placing Plesiastraea spongiformis in the synonym of 
Stephanoeoenia intersepta. 

Plesiastraea goodei Verrill, fragment of the type No. 36497, 
U.S.N.M., is precisely the same as Stephanoeoenia intersepta — there 
are no differential characters. 

Distribution of Stephanoeoenia intersepta. — Just how old, geolog- 
ically, this species is, is not definitely known. 
* Jamaica. — There is a specimen in the United States National 
Museimi bearing the station nimiber 2580, which is for the collection 
made by Messrs. J. B. Henderson and C. T. Simpson in the Bowden 
marl of Jamaica. 

Santo Domingo. — ^Miss C. J. Maury obtained five specimens of this 
. much-named species, as follows: 

Rio Gurabo: Zone D, associated with Stylophora affinis Dimcan, 
Madracis decaetis (Lyman), PoetUopora crassoramosa Duncan, OrhiceHa 
limhata (Duncan), OrhiceUa cavernosa var. cylindrica (Dimcan), iSy^y- 
gophyUia dentata (Duncan); zone E, associated with Placoeyathus 
new species, Placoeyathus variabilis Dunc^m, Stylophora new species, 
Madracis decadis (Ljrman), SyzygophyUia dentata (Dimcan), Pavona 
new species. Limestone, Los Quemados, associated with Placoey- 
athus variabilis Duncan. As zones I and H of Miss Maury's sec- 
tion repiMent the Bowden fauna, zones E and D are stratigraphi- 
cally above the Bowden. 

Ouha. — I collected a specimen near the Morro, at the mouth of 
Santiago Harbor, altitude about 240 feet above level. This specimen 
may be of Pleistocene age. The general basement country rock is 

3714»— 19--Bull. 108 12 



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360 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Miocene limestone and marl, which contain some corals of reef fades ; 
and on this basement there are in places well-developed Pleistocene 
coral reefs. Therefore, the specimens of Stephanocoenia irUersepUi 
might be of Miocene age. Other specimens from stations 3436 and 
3449, south side of the trocha in Santiago, seem definitely to belong 
in the La Cruz marl and to be of pre-Pleistocene age. 

Doctor Pittier obtained a specimen of the species at the ''CoUine 
en d^moUtion/' Limon, Costa Rica, apparently in association with 
AsterosmUia hiUi, Dichocoenia ivherosa, and BaianophyUia pittieri. 
The horizon would therefore be near that of the Bowden marl. 

Pleistocene. — Gteneral in the elevated reefs of the Caribbean and 
Gulf region: Barbados (low-level reefs); Curasao and Arube; Key 
Vaca, Florida. 

Recent. — ^The West Indies in general, northward to the Bermudas; 
Florida; British Honduras. 

Although I have often picked up specimens of this species where 
they had been washed up by the waves, both in Florida and in the 
Bahamas, I have not certainly seen it alive on the reefs. As the 
color of the living polyps is brown, while alive it so closely resembles 
Siderastrea rctdians that only very close examination will distinguish 
between them, probably on the reefs it was mistaken for the latter. 
That it is a common associate of the usual West Indian reef corals 
is shown by its usual presence among them in the fossil reefs. This 
species ranges into slightly deeper water than most of the West 
Indian reef corals. I dredged-it at a depth of 4-9 fathoms oflf Nas- 
sau, Bahamas, and at a depth of 16 fathoms oflF Tortugas, Florida. 
Genus DICHOCOENIA MUne Edwards. 

1848. Dichocoenia Milne Edwards and Haiue, Compt. Rend., vol. 27, p. 469. 
1857. Dichocoenia Milne Edwards and Haime, Hist. nat. Corall., vol. 2, p. 199 

(type-species, figured, pi. DI, figs. 10a, 106).* 
1917. Dichocoenia Vaughan, U. S. Geol. Surv. Prof. Pap. 98-T, p. 370. 

Type species, — Dichocoenia porcata Milne Edwards and Haime. 

DICHOCOENIA TVBEROSA Danean. 

Plate 79, figs. 4, 4a, 4b, 

1863. Dichocoenia ttiberosa Duncan, Geol. Soc. London Quart. Joum., vol. 19, 
p. 432, pi. 15, figs. 5a, bb. 

This name has been placed in the synonymy of the living Dicho- 
coenia stoJcesi Mihie Edwards and Haime by both Gregory * and my- 
self.' One-half of Duncan's type is in the United States National 
Musemn, No. 155275, presented by the officers of the Geological 
Society of London. Although D. tuberosa is very similar to D. 
stokesif D, tuberosa has a pendimculate base and granulate costal 
markings below the calicular surfaces in all the specimens I have 

1 Oool. Soc. London Quart. Joum., vol. 51, p. 268, 1886. 
« U. 8. 0«ol. Survey Prof. Pap. 98-T, p. S71, 1017. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 361 

examined. As I am able to recognize the species I am treating it 
as valid. Dimcan records the form from the '^Nivajd shale and 
tufaceous limestone of Santo Domingo." 

Locality and geologic occfwrrence. — Costa Kica, ''Colline, en demoli- 
tion/' Limon, No. 618 of H. Pittier collection, associated with 
Asterosmiliahmij StepJianocoeniaintersepta, and BcdanophyUia pjMieri. 
A single, small, immatm*e specimen. The illustrations present its 
characters well enough to make a detailed description minecessary. 

Santo Domingo, Rio Gurabo, zone F, of Miss C. J. Mamy's sec- 
tion, associated with Placocxjathus variabilis Dmican and AntiUia , 
iluhia (Duncan). 

Genus EUSMILIA Milne Edwards and Haime. 

1848. Eusmilia Milne Edwards and Haime, Comptes Rend., vol. 27, p. 467. 
Type-species. — Madrepora fastigiaia Pallas. 

EUSMIUA FASTIGIATA (PiOlaB). 

1 7 66 . Madrepora fastigiata Pa llas , Elench . Zooph . , p . 301 . 

1895. Eusmilia fasHgiata Gregory, Geol. Soc. London Quart. Joum., vol. 51, 

p. 260 (with synonymy). 
1895. Euxmilia knorri Gregory, Geol. Soc. London Quart. Joum., vol. 51, p. 261 

(with synonymy). 

1901. Evsmilia knorri Vauohan, Geol. Reichs. Mus. Leiden Samml., ser. 2, vol. 

2, p. 13. 

1902. Eusmilia aspera Verrill, Conn. Acad. Arts and Sci. Trans., vol. 11, p. 114, 

fig. 3. 

1915. Eusmilia fasHgiata Vauohan, Washington Acad. Sci. Joum., vol. 5, p. 596. 

1916. Eusmilia fastigiataY AVQKAV, Carnegie Inst. Washington Yearbook No. 14, 

p. 227. 

Study of large suites, of Eusmilia convince me that Eusmilia 
fastigicUa (Pallas) and E. aspera (Dana) = E, Jcnorri M. Edwards and 
Haime are not specially separable, as there is great variation and 
complete overlapping in the columellar characters by which they 
were distinguished. 

Localities and geologic occurrence. — Canal Zone, Pleistocene at sta- 
tions 5849, Mount Hope; Costa Rica, 6251, Monkey Point, collected 
by D. F. MaoDonald. 

General in the living and Pleistocene coral reefs of Florida, the 
West Indies, and the Caribbean coast of Central America. 

Family ASTRANGHD^ Verrill. 
Genus CLADOCORA Ehrenberg. 

1834. Cladocora Ehrekbero, Gorallenth. Roth. Meer., p. 85 (of separate). 
1848. Cladocora Uome Edwards and Haimb, Comptes Rend., vol. 27, p. 493. 

Typ&'Species. — CaryophyUia cespitosa Lamarck. 



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362 BriJ.ETIN 103, UNITED STATES NATIONAL MUSEUM. 

CLADOCORA ARBUSCULA (Le Suev). 

1820. Caryopkyllia arbuscula Le Sueur, Paris Mus. M^m., vol. 6, p. 275, pi. 15, 

figs. 2a-2rf. 
1901. Cladocora arbuscida Vauohan, U. S. Fiah CommisBion Bull, for 1900, vol 2, 
p. 298, pi. 2, figB. 3, 3a (with synonymy). 
This species is common in the Pleistocene marls near Colon. 
Locality and geologic occurrence, — Canal 2iOne, station 5850 and 
6039, Pleistocene, Mount Hope, collected by D. F. MacDonald. 
Living in Florida and the West Indies on reef flats and in water from 
8 or 9 to about 20 fathoms deep. 

Family ORBICELLIDAE Vaughan. 
Genus ORBICELLA Dana. 

1846. Orbicella Dana, U. S. Expl. Exped. Zooph., p. 205. 

1849. Phyllocoenia Milne Edwards and Haiub, Gomptes Rend., vol. 27, p. 4^^ 

1901. Orbicella Vaughan, Geol. Reichs Mus. Leiden Samml., ser. 2, vol. 2, p. 21 

1902. Orbicella Verrill, Conn. Acad. Arts and Sci. Trans., vol. 11, p. 93. 
1918. Orbicella Vaughan, Camegie Inst. Washington Pub. 213, p. 85. 

Type-species, — Mcbdrepora annularis Ellis and Solander. 

Of this perplexing genus of corals, the following species and varie- 
ties are treated as valid in the present papers: 
Orbicella annularis (Ellis and Solander), 
lirnbata (Duncan). 
imper^UoriSy new species. 
aUissima (Duncan). 
aniiUarum (Duncan). 
ca/vemosa (Linnaeus). 

var. endoihecata (Duncan), 
var. cylindrica (Duncan). 
aperta (Verrill). 
bainhridgensiSf new species. 
cosiata (Dimcan). 
canaKSf new species. 
tampdensis, new species. 

var, sUecensis, new variety 
brevis (Duncan). 
insignis (Duncan). 
intermedia (Duncan). 
gdbbi, new species. 
As synonymy is discussed on subsequent pages, it is here only 
necessary to say that under the name Astra^ea megalaxona ^ Duncan 
described from Antigua a silicified coral which is not determinable; 
that his Astraea crassolameUMa ' and its varieties are here referred to 

> (Jeol. Soc. London Quart. Journ., vol. 19, p. 420, pi. 13, flgs. 128, 126, 1883. 
« Idem., p. 412, pi. 13, fiRs. 1-7. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 363 

trlxe fungid genus Difloastrea Matthai; his Aatraea ceUulosa ^ is made 
tl\e type-species of a new genus, An^guastreaj and his Aatrcbea anti" 
^t^ensis^ and Astraea tenuis^ are referred to the fungid genus 
C^cUhcmoTTpha Reuss. 

Although inadequacy of information regarding four species, O. 
aJtissima, 0, andUarumj 0. insigniSj and 0. intermediaj described by 
Ouncan, renders the preparation of an adequate synoptic tabk 
impracticable, an attempt will be made to summarize the most 
striking characters. With one exception, the species fall into two 
l&rger groups: the members of the first group normally have only 
tliree cycles of septa; those of the second group have four cycles, 
the fourth cycle is incomplete in some specimens, while in other spec- 
imens a variable mmiber of quinary septa arc present. Orlo species^ 
OrbiceUa gabhi Vaughan, has five cycles of septa. 

SYNOPSIS OP AMERICAN SPECIES OF ORBICELLA. 

Species with S cycles of septa. 

Oalicee usually 2 to 3 mm. in diameter; costae subequal; primary and secondary septa 
equal, extend to the columella 1.0. annularis (Ellis and Solander). 

Calicefl 3 to 4 mm. in diameter; costae usually alternately large ftnd small; secondary 
septa thinner than the primaries, but usually reach the columella 

2. 0. limhata (Duncan). 

Calicefl 3.6 to 5 mm. in diameter; costae prominent, thin; secondary septa usually 
about half as long as the primaries, tertiariee small and thin. 

3. 0. imperatorisj new speciea. 

Calices 7.5 mm. in diameter; costae tolerably developed, subequal; primary and sec- 
ondary septa subequal, extend to the columella... 4. 0. antillarum (Duncan). 

Species of OrbiceUa with the 4th cycle of septa nearly or quite complete. 

Calicee 5 mm. in diameter; costae unequal, thicker than the septa, last ^'order!*' of 

costae well developed, contrasting with rudimentary septa; septa irregular in 

arrangement, 36 in number, 6 septa in each of 6 systems. . b.O. altissirna (Duncan). 

Calices from 5 to 11 mm. in diameter; costae correspond to all septa, usually subequal; 

septa normally in 4 complete cycles, subequal over top of wall, first 3 cycles reach 

columella, no pali 6.0. cavernosa (Linnaeiis). 

Gestae strongly alternating in size, fourth cycle small and thin without ob- 
vious corresponding septa 6a. var. endothecata (Duncan) . 

Corallites smaller than in 6a (5 to 6 mm. in diameter), about. 38 septa, last 

cycle of costae rudimentary or obsolete 66. var. cylindrica (Duncan). 

Similar to 0. cavernosa except that the first three cycles of septa are thinner and 
taller, strongly contrast in height with the quaternaries. . 7.0. aperta (Verrill). 
Calices 6 to 7 mm. in diameter; costae low, equal; septa low and subequal on mural 
summit; primaries and secondaries with rather wide erect, paliform lobes, young- 
est septa composed of incompletely fused spines. 

8. 0. bainbridgeJisiSj new species. 
Calices 7.5 to 8.5 mm. in diameter; costae highly developed, alternate in size except 
at calicular margin; septa normally in 4 cycles, thin except in wall of some speci- 
mens, paliform lobes and thickenings distinct but rather small, tertiaries usually 
shorter than secondaries. 9. 0. costata (Duncan). 

1 Qeol. Soc. London Quart. Joom., vol. 10, p. 417, pi. 13, flg. 10. < Idem. p. 419, pi. 13. fig. H. 
sidem,p.421,pl. 13, flff. H. 



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364 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Oalices 5 to 9 mm. in diameter; costae subequal or alternately large and anudl belor 
calicular edge. Septa in 4 or nearly 4 complete cycles; primaries as a rule notably 
laiger than the secondaries, with a prominent tooth on inner end; secondaries 
smaller, but with paliform tooth on inner end of each; tertiaries still smaller: 

quaternaries very small 10. 0. eanalis, new spedee. 

Calicos 6 to 10 mm. in diameter, exsert 4 to 4.5 mm.; costae very prominent, no or 

only rudimentary costae correspond to last cycle of septa; septa in 3 or 4 sizci. 

maiginsof primaries exsert as much as 1.5 mm. . : 11. 0. Uanpaennu, new spedaF. 

Calicos not so elevated as in 11; small but distinct costae correspond to lasi 

cycle of septa 11a. var. tUecensis^ new varkty. 

Calicos 5 mm. in diameter, protuberant but rather low; costae strongly alternating 
in size; primary septa the largest; fourth cycle incomplete 

12. 0. 6rmf (Duncan . 

Calicos 10 mm. in diameter; costae long, slender, subequal, occasionally a rudim^itan 

costa with no corresponding septum; septa delicate, long, slender, distant, fourth 

cycle incomplete 13. 0. insignis (Ehincan . 

Calicos 5 mm. in diameter; in places small costae between larger ones; a few qua- 
ternary sei^ta 14. 0. intermedia (Duncan; 

The numbers preceding the names in the synopsis correspond to 
numbers before the names heading the following descriptions. 

As OrhiceUa gahbi is the only species with 5 complete cycle of septa, 
it needs no special caption nor is 0. irradians included in the key. 

1. OBBICELLA ANNULARIS (Snis ud S«kuMler). 

Plate 80, figs. 7, 7a, 76; plate 81, figs. 1, la, 2; plate 82, figs. 1, la, 2; plate 83, ^. 
1, 2, 3, 3a,- plate 84, figp. 1, 2, 3, 3a. 
1786. Madrepora annularis Eixis and Solander, Nat. Hist. Zooph., p. 169, pi. 53, 

figs. 1, 2. 
1786. Madrepora faveokUa Ellis and Solandbb, Nat. Hist. Zooph., p. 166, pLoS, 

figs. 5, 6. 
1790. Madrepora acropora Gmblin, Linn. Syst. Nat., ed. 13, p. 3767. 
1790. Madrepora faveolata Gmeun, Linn. Syst. Nat., ed. 13, p. 3769. 
1794. Madrepora acropora Esper, Pflanzenth., Fortsetz., vol. 1, p. 21, pi. 38. 
1816. Astrea annularis Lamarck, Hist. nat. Anim. s. Vert., vol. 2, p. 259. 
1821. Astrea annularis Lamouboux, Exp. M^th. Genres dee Polyp., p. 58, pi. 53, 

figs. 1, 2. 
1821. Astrea faveolata Lamouboux, Exp. M^th. Genres des Polyp., p. 58, pi. 5S, 

figs. 5, 6. 
1834. Ezplanaria annularis Ehrbnberq, Gorallenth. Roth. Meer., p. 84 (of 

separate). 
1846. Asthiea (Orbicella) annularis Dana, U. S. Expl. Exp. Zoophytes, p. 214. 

pi. 10, fig. 6. 
1857. Heliastraea annularis Mn<NB Edwards and Hadcb, Hist. nat. Corall., vol. 

2, p. 473. 
1861. Heliastraea annularis Duchassaing and Michblotti, M6m. OoraU. AntiUes, 

p. 76 (of reprint). 
1861. Heliastraea acropora Duohassaino and Michblotti, M6m. (3oraIl. Antilles, 

p. 76 (of reprint). 
1861. Heliastraea lamarcki Duchassaing and MiCHELom, M6m. Corall. Antilles, 

p. 76 (of reprint). 
1863. Cyphastraea costata (part) Duncan, Geol. Soc. I>ond. Quart. Joum.. vol. 19, 

pp. 441 and 443. 
1863. Astraea harbadensis Duncan, Geol. Soc. Lend. Quart. Joum., vol. 19, pp- 

421 and 444, pi. 15, figs. 6a, 66. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE. 365 

1864. Orhicella anntUaris Vbrrill, Mus. Ck)mp. Zool. Bull., vol. 1, No. 3, p. 48. 

1865. Orbicella anntdaTis Verrill, Boston Soc. Nat. Hist. Proc., vol. 10, p. 323. 

1866. Heliastraea annularis Duchassainq and Michelotti, Sup. Mto. Corall. 

Antilles, p. 84 (of reprint). 
1866. Heliastraea lamareki Duchassatng and Michelotti, Sup. M^m. Corall. 

Antilles, p. 84 (of reprint). 
1866. Heliastraea aeropora Duchassaino and Miohblotti, Sup. Mdm. Corall. 

Antilles, p. 84 (of reprint). 
1866. Heliastraea barbadensis Duchassaino and Michelotti, Sup. Mto. Corall. 

Antilles, p. 85 (of reprint). 
1866. Cyphastraea costata Duchassaino and Michelotti, Sup. M^m. Corall. 

Antilles, p. 85 (of reprint). 
1868. Heliastraea barbadensis Duncan, Geol. Soc. Lond. Quart. Joum., vol. 24, 

p. 24. 
1868. Cyphastraea costata Duncan, Geol. Soc. Lond. Quart. Joum., vol. 24, p. 24. 
1895. Orbicella aeropora Gregory, Geol. Soc. Lond. Quart. Joum., vol. 51, p. 272. 
1895. Cyphastraea costata Gregory, Geol. Soc. Lond. Quart. Joum., vol. 51, p. 274. 
1895. J^nopora/rarwH Gregory, Geol. Soc. Lond. Quart. Joum., vol. 51, p. 274, 

pi. 11, figs. 2a, 26. 
1901. OrbioeUa aeropora Yaughan, Geolog. Reichs. Mus. Leiden Samml., ser. 2, 

vol. 2, p. 22. 

1901. Orbicella aeropora Vaughan, U. S. Fish Commission Bull, for 1900, vol. 

2, p. 301, pis. 6, 8. 

1902. Orbicella annularis Verrill, Conn. Acad. Arts and Sci. Trans., vol. 11, 

p. 94, pi. 15, ^, 1. 
1902. Orbicella annularis var. stellulata Verrill, Conn. Acad. Arts and Sci. 

Trans., vol. 11, p. 96, pi. 15, fig. 2. 
1902. Orbicella hispidula Verrill, Conn. Acad. Arts and Sci. Trans., vol. 11, p. 

100, pi. 15, figs. 3, 3a, 36. 

1902. Orbicella annularis Vaughan, Biol. Soc. Washington Proc., vol. 15, p. 56. 

1903. Orbicella annularis Duerden, Nat. Acad. Sci. Mem., vol. 8, p. 564, pis. 

8-10, figs. 64-73. 

1915. Orbicella annularis Vaughan, Washington Acad. Sci. Joum., vol. 5, p. 596. 

1916. Orbicella annularis Vaughan, Camegie Inst. Washington Yearbook No. 14, 

p. 227. 

Subsequent study has led me to believe that changes should be 
made in the synonymy as given in the first of my papers cited in the 
synonymy. PhyUocoenia limhata Duncan, P, limhata var. tegula 
Duncan, and Plesiastraea ramea Duncan represent one species and it 
is separable from OrbiceUa annularis. As PhyUocoenia limhata is the 
older name, the species should be designated Orhicella limhata (Dun- 
can). The most conspicuous difference between it and 0, annularis 
consists in its primary septa being markedly more developed than 
the secondaries. 

Orhicella annularis is the principal coral of the outer reefs in Florida, 
the West Indies, and on the Caribbean side of Central America. Jt 
is general in the elevated Pleistocene of the same region. 

Prof. J. Graham Kerr, of the University of Glasgow, has kindly 
sent me photographs of the type of this species, which is preserved 



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366 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

in the Hunterian Museum at that institution, and I have based the 
following description on them: 

The corallum is head-shaped, with a greater diameter of 107 mm. 
and a lesser of 86. 

The calices are circular, 2 mm. in diameter, margins slightly ele- 
vated, joined by equal costae, distance apart usually about 1 mm., 
occasionally 2. 

Septa 24 in number, alternately larger and smaller; the lai^r are 
rather thick and reach the columella; the intermediate ones are short 
and their inner ends are free. 

Columella spongy, well developed, its diameter about one-third 
that of the calice. 

A comparison of the photographs with specimens shows that the 
traditional OrbiceUa anTwlaris of the Caribbean and Gulf region is 
correctly identified. 

There are in the collection of the United States National Museum 
a number of specimens that are almost duplicates of the type-speci- 
men, except that they are not worn, as is the type. These specimens 
form the basis of the succeeding description (see pi. 81, figs. 1, la). 

The corallum forms rounded masses rising above a rather lai^, 
firmly attached base, which is, however, less in diameter than the 
maximum diameter of the corallum. Frequently there is a pro- 
jecting or incrusting edge whose lower surface is covered by epitheca. 
The upper surface may be imiformly roimded, undulate, or lobed. 
The size, of course, is variable; the masses may be several feet in 
diameter. 

The caUces are circular, or slightly deformed. Their diameter, 
measured betweeh thecal smnmits is from 2 to 2.6 man. In depres- 
sions oD the surface they may be smaller, about 1.5 mm., but these 
are abnormal. Their edges are from 0.5 to almost 2 mim. apart, 
about 1 mm. is probably an average. The calicular edges are 
slightly elevated. The intercorallite areas are costate. Costae cor- 
respond to all septa; subequal or alternating in size, those of adjoin- 
ing calices meeting; edges dentate; thicker than the width of the 
intercostal spaces and moderately elevated. 

Septa in three complete cycles, primaries and secondaries equal, 
rather stout, extending to the columella and fusing to it; tertiaries 
shorter, about half the length of the primaries, somewhat thinner, 
inner edges free. Margins of the primaries and secondaries dend- 
edly exsert; their inner edges fall perpendicularly to the bottom of 
the calicular fossa, and bear just above the columella one or two 
prominent teeth, with a few smaller teeth above; the septal arch is 
either very gentle, obtuse, or it may be tnmcate, its dentations fine; 
the outer margins steep, but mor(B inclined than the inner, dentations 
relatively coarse. Septal faces finely granulate; in longitudinal 
actions, the inner edges are lacerate, the last cycle with perforations. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 367 

Gndothecal dissepiments delicate, thin, nearly horizontal, shghtly 
inclined downward from the corallite wall8> In this series of speci- 
inens the corallite walls are thick and close together, those of ad- 
jacent corallies sometimes being solidly fused together; usually, 
lio-wever, there is some exotheoa, consisting of stout, subhorizontal 
dissepiments. 

Columella well developed, formed by interlacing processes from 
-the inner edges of the septa; diameter from one-third to one-half 
that of the calice; its upper surfcce about 1 mm. below the thecal 
xnargin. 

These specimens, it should be repeated, are typical, and except in 
size and to a certain extent in the configuration of the surface show 
abnost no variation. They come from the following localities: Dry 
Tortugas, Florida, Dr. Edward Palmer, collector, 8 specimens; east 
end of Hog Island, Bahamas, B. A. Bean, collector, 1 specimen; 
Florida and the Bahamas, many specimens, collected by T. W. 
Vaughan and others. There are other specimens, bearing the in- 
definite label ''West Indies" or having no locality stated. These 
localities indicate that the species in its typical foim is of general 
occurrence in the coral reef areas around the Caribbean Sea and Gulf 
of Mexico. 

The recent specimens in the United States National Museum show 
at least four kinds of variation from the typical form. 

Variation No. 1 (pi. 84, fig. 2J. — ^This variation is, I believe, only 
a growth form. It, in its structural features, is the same as the 
typical form, except that the septa near the growing edge are less 
exsert and the exotheca appears to be absolutely solid. The corallum 
is an obtuse, comja^essed colunrn, with an undulated surface. Greater 
diameter of the base, 62 mm.; lesser 52 nun.; height 72 mm. 
Locality.— Dry Tortugas, Florida. 

Variation No. 2 (pi. 81, fig. 2).— The general growth form is similar 
to that of typical specimens, except that the surface is thrown iilto 
gibbosities of irregular shape and size; these are often about a centi- 
meter in height and several centimeters in diameter. The calices 
are laj^er than in the typidkl specimens, often measiuing 3, occa- 
««ionally 4 millimeters in diameter, between thecal summits. The 
thecal edges are slightly elevated; the margins of the primaries and 
secondaries decidedly exsert, not infrequently standing 2 mm. above 
the intercorallite furrow. The three characters here mentioned are 
the distinguishing ones of this variation, namely, gibbosities on the 
surface; larger calices; and more exsert septa. 

Localities. — Dry Tortugas, Florida, Dr. Edward Palmer, collector, 
1 specimen; east end of Hog Island, Bahamas, B. A. Bean, collector. 
1 specimen; and two other specimens, without locality labels. 



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368 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

Variation No. S (pi. 82, fig. 2) is represented by a single specimen. 
The corallum is discoid, lower surface flat, upper surface convex, some 
irregularities. Greater diameter, 22.7 cm., lesser, 19.2 cm; thicknffy 
in the center about 5 cm., on the edge, 3 cnu 

Calices with elevated margins and crowded together, the different 
corallite walls almost contiguous; margins of primary and secondary 
septa decidedly exsert. Diameter of caUcos about 2.75 mm. 

The distinguishing charact^ of this variation are its discoid f<»iu, 
its crowded caUces, its decidedly exsert septal margins. 

Locality. — Fort Taylor, Key West, Florida. 

Variation No. 4 (pl- 82, figs. 1, la) is represented by the speci- 
mens that I have described from Mayaguez, Porto Kico, in my 
''Stony corals of the Porto Rican waters." * The following descrip- 
tion is based on them: • 

The corallum forms ascending masses; the largest specimen i> 
about 20 cm. tall; diameter above flawed -out base about 13.5 cm. 
The base of each specimen is considerably produced as a wide, free 
edge invested below by epitheca. 

Calices with very slightly or only moderately elevated margins, 
diameter measured between Qxecal summits, from 3.25 to 4 muL: 
rather shallow; distance apart, from a thin dividing edge to 2.5 
mm.; about 1.5 mm. is probably the average. Thin costae moder- 
ately prominent, subequal, or aJtemating in size, correspond to all 
septa; those from one calice extend across the intercorallite spaces 
and meet those from the adjacent calices. 

Septa thin, 24 to 28 in number, one-half of them extend from the 
wall to the columella, and have decidedly exsert margins; the other 
half are not so tall and are short, their inner ends free. 

Endotheca and exotheca as in the typical specimens, except tiiat 
they are more delicate. 

These differ from typical specimens by their much lighter texture, 
which, of course, is determined by their thinner skeletal structures, 
the wide, flaring, free edges of the base, and their lai^r calices. 
The caUces overlap in size those of variation No. 2, otherwise 1 
should consider the specimens as representing a distinct species. 

Variation No. 6 (pi. 83, figs. 1, 3, 3a). — OrhiceOa hiapidvla Verrill.' 
The following is the original description: 

Coral an incrusting mass over 125 mm. across, and from 5 to 20 mm. ^ck. The 
texture is rather solid and heavy, there being much solid exotheca between the 
calicles, which are rather far apart, the interspaces being mostly equal to, and often 
exceeding, their diameter. 

The calicles are round, regularly stellate, a little prominent, with swollen, sloping, 
costate rims much as in those of 0. annularis, which they -resemble in size, thougb 
distinctly larger. The septa are in three very regular cycles; the twelve principal 

1 Bull. U. S. Fish Commission for 1900, vol. 2, p. 301, pb. 6, 7, 1901. 

s Conn. Acad. Arts and Sd. Trans., vol. 11, pp. 100, pi. 15, figs. 3, So, 36. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 369 

ones are wide, nearly equal, all reaching the rather large columella; their edges are 
perpendicular and finely, sharply serrate, with slender, rough teeth, which extend 
also over their prominent, obtuse, or subtruncate summits, giving them a rough 
ap]$earance under a lens; their surfaces are also rough or hispid with numerous conical 
Strains. The septa of the third cycle are narrow, straight, and usually reach about 
lialiway to the columella. 

T!-e coetae are thick, not very high, meeting or inosculating between the calicles, 
and covered with a single row of small slender, rough spinules. The colimiella is 
\^e\\ developed, formed of contorted trabecular processes, and often having a small 
pit in the center and a few erect spinules, similar to the slender, rough, paHform teeth 
that often (but not regularly) stand at the base of some of the 12 larger septa. 

In sections the walls are very thick and nearly solid. The endothecal dissepi- 
ments are small, thin, irregularly convex or flat above. The calicles are not filled 
lip below, or only slightly encroached upon, by a deposit between some of the septa. 
IMameter of the calicles 3 to 3.5 mm.; distance between them mostly 2 to 4 mm., 
often more. 

Florida Reefs (Maj. E. B. Hunt), Yale Museum, Xo. 98. Near Nassau, N. P. 
(coll. R. P. Whitfield), Amer. Mus., New Y'ork. 

This has the general appearance of 0. annularis, but with calicles larger than usual 
and decidedly farther apart. The walls and exotheca are much thicker and more 
solid, and the endothecal cells are fewer and less regular. The sharply spinulose 
and hispid septa and coetae are also characteristic. The exothecal deposits are 
nearly as solid as in Oculina. 

A Nassau specimen, in the American Museum, is an irregular, rounded mass about 
5 inches in diameter, and 3 to 4 thick, with a a lobulated surface. The coral is heavy 
and solid; the surface of the coenenchyma is spinulose; the coetae well developed. 
The calicles are more variable in size than in the type, in some places being one-half 
emaller and closely crowded. Goll. R. P. Whitfield. 

The form of 0. Tiispidula Verrill, in which the upper surface is 
lobulate, is common on the reef off Cocoanut Point, Andros Island, 
Bahamas, where a suite of 12 specimens was obtained by the Anton 
Dohm expedition in 1914. The calices of most of these specimens 
are precisely as in the type of Professor Verriirs 0. hispidula (frag- 
ment of type No. 40476, U.S.N.M.) and Gregory's Echinopora franslci 
(fragment of type No. 156455, U.S.N.M.), but in both growth 
form and calicular characters there is intergradation with the more 
usual characters of 0. annularis. Plate 76, figures 3, 3a illustrates 
the appearance of one of the specimens with lobulate surface. 

A specimen from Port Castries, Santa Lucia (pi. 83, fig. 2), shows 
a variation worthy of note. In all of the variations so far described, 
the primary and secondary septa are constantly subequal, uniformly 
reaching the coliunella. In the Santa Lucia specimen a secondary 
septum in some systems is shorter and thinner than a primary; and 
in some calices there are as many as 30 septa. This specimen is of 
importance for comparison with PhyUocoenia scvZpia var. tegula 
Duncan and Echinopora fravJcsi Gregory. 

These remarks cover the variation of the recent specimens that 
I have actually been able to study. Pourtalte, Verrill, and Duerden, 
however, have added other observations. 



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370 BULLET!?^ 103, UNITED STATES NATIONAL MUSEUM. 

Pourtalds says of the species: 

The same remarks about variatdon, given under the head of 0. cavemoMy can be 
applied to this species; there are very fine examples in the museum of the great 
variation of form of the caUcles in the same specimen. 

It is very common in Florida on the reef and in the channels, and forms large 
hemispherical masses nearly up to low-water mark. The central and highest part 
often dies out from being left uncovered at very low tide and the mass then assumes 
an annular form through the decay of the dead part.* 

Veriill writes: 

It shows considerable variation in the size of the calicles; in the extent to which 
they are crowded together; in the prominence of their borders above the intervening 
exotheca; in the prominence of the septa above the walls; and in the extent to whidi 
the small septa of the third cycle are developed. But yet these variations, so far as 
I have seen, never go so far as to render difficult the recognition of the species unless 
the specimens are badly worn. 

****** • 

When well grown it forms hemispherical or spheroidal masses, up to 5 feet or more 
in diameter. But it also grows in irregular incrusting plates, and sometimes in nodose 
or lobulate masses, or even in branched forms .^ 

Duerden, in describing specimens from Jamaica, says: 

The species occurs on coral areas in small or large, fixed, nearly spheroidal masses; 
also as an incrustation occupying areas several feet across. Small isolated colonies 
are sometimes conical. . In places it is an important constituent of the reefs.' 

This is one of the species to which I devoted a great deal 
attention in my study of the living reefs in Florida and the Bahamas, 
and have inserted references to two of my papers (1915, 1916) in which 
it is considered. It is preeminently the great reef-building species 
of the Pleistocene and Recent reefs in Florida and the West Indies. 
Where there is sand on the bottom, it forms tall, thick, round-topped 
columns. 

VARIATION OF FOSSIL SPECIMENS. 

There are specimens, particularly those of known Pleistocene age, 
similar to the typical form of the species, except that there may be 
variations in the size of the calices; those of a specimen from Fort 
Nassau, Curavao, range from 3 to 4.6 mm. in diameter, measured 
between thecal summits; those of another specimen from West- 
punt, Curavlao, are from 2.5 to 3 mm. in diameter. The former 
possesses the largest calices of any specimen of the species I have seen. 

The variations not included in the preceding remarks may be 
divided into two classes, dependent upon growth — namely, a, ex- 
planate or incrusting; 6, colmnnar. 

A. Growth from cxplanate or incrusting. 

1 Ma& Comp. Zool. lU. Cat. No. 4, p. 72, 1871. 

« Conn. Acad. Arts and Sci. Trans., vol. 11, pp. 05, 96, 1902. 

* West Indian Madrepnrarian Polyps. Nat. Acad. 8d. Mem., vol. », p. Ml, 1908. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 



371 



Gregory* was mistaken in referring the specimens described by 
him as Echino'pora franlcsi (see pi. 84, fig. 4) to the genus Echinopora. 
The following is the original description: 

Diagnosis, — ^The coral has a broad base; from this pass outward short, thick, rapidly- 
tapering expansions. 

Corallites long, often an inch in length. Their distance one from the other varies 
from half their diameter to the whole. 

Septa strongly dentate; inner teeth paliform, in three cycles. Those of the first 
cycle always unite to the columella; those of the second cycle often do so, but may 
join the primary septa; those of the third cycle are much smaller and independent, 
but a few may unite with the septa of the other orders. 

Columella of very loose tissue; half the diameter of the corallite. Endotheca scanty. 
Ooenenchyma thinner than in other species of the genus. Echinulations of the surface 
coarse. Epitheca thick and well developed. 

Dimensions. — Diameter of an average corallite, 3 mm.; height of corallite varies 
from 10 to to 25 mm.; thickness of wall varies from 1) to 3 mm. 

DistnbiUion. Recent: West Indies. Fossil: Barbados: Lowlevel Reefs, near 
Bridgetown. 

Cotypes, — British Museum (Natural ffistory); a piece of one of 
the cotypes in the United States National Museum, No. 156,455. 

A comparison of this description with the notes on the variation 
of OrhiceUa annularis will show that it presents no important differ- 
ence from- variations of the species already recorded. Its growth 
form is explanate, the exotheca is solid, and the secondary septa 
often, but hot always, reach the colimiella. 

B. Growth from columnar (pi. 84, figs. 3, 3a.) 

These are the specimens referred to in my paper "Some fossil 
corals from the elevated reefs of Curasao, Axube, and Bonaire, '* ^ 
obtained by Mr. v. Koolwijk at Westpunt, Curasao. Three of the 
specimens are in the United States. National Museum, and they 
form the basis of the following description : 

The coralliun forms ascending, compressed, obtuse columns. 
Dimensions in millwuters of variant o/OrHcella annularis from Curasao. 



Specimen No. 


Qieat«r 
diameter 
of base. 


Lesser 
dismeter 
or base. 


1 


mm. 
37.5 
30+ 

27.6 


mm. 
35 
23 

21 


2 


3 





Height. 



Remarks. 



mm. I 

60 I BJftiroatlon 82 mm. above base. 

71 Constricted above base; gradually enlarging 

above the constriction. 
91 Figured, pi. 84, figs. 3, 3a. 



Calices 2.5 to 3.5 mm. in diameter; from less than 1 mm. to 2 nmi. 
apart. • The upper margin is usually not elevated, while the lower 
one is, tiius tilting the calicular orifices. The maximum length of 
the lower limb of the calice is about 3 mm. Subequal, relatively 
thick, dentate costae correspond to all septa. 

> Oeol. Soc London Quart. Jour., vol. 51, p. 274, pi. 11, figs. 2a, 25, 1805. 

> Qeolog. Reichs. Mus. Leiden Samml., ser. 2, voL 2, Heft I, p. 26. 



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372 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

The usual number of septa is three complete cycles; primaries and 
secondaries subequal, reach the columella; tertiaries short, inner 
edges free. The septa present only one noteworthy difference from 
what is usual in 0. anrmlaria; that is, the margins of the primaries 
and secondaries are less exsert. 

Columella not very large, loose, trabecular. 

The three saUent characteristics of this variant are (1), its growth 
form; (2), the tilted caUces; (3), the lower (\ess exsert) margins of 
the primary and secondary septa. 

Geologic horizon. — Pleistocene. 

NOTES ON SYNONYMY. 

A number of other names need to be considered in greater or less 
detail. 

Gr^ory ^ apphed the name OrbiceUa acropora (Linnaeus) to this 
species. He accepted the determination of the species by Milne 
Edwards and Haime,* who separated it from 0. annularis by its 
having no septa corresponding to the last cycle of costae. Gr^ory 
showed that occasionally in typical specimens of 0, annularis the 
last cycle of septa may be absent while the costae are present, thus 
brealdng down the character used by Milne Edwards and Haime to 
distinguish the species. I accepted Gregory's conclusion, and 
followed him in my paper on Some fossil corals from the elevated 
reefs of Curasao, Arube, and Bonaire, and subsequent papers. 
Professor VerrUl, in his Variations and Nomenclature of Bermudian, 
West Indian, and Brazilian reef corals,' declares that Madrepora 
a^cropora Linnaeus *'is utterly indeterminable," and takes the next 
later specific name, annularis Ellis and Solander, for the species. 
Subsequent study convinced me that Professor Verrill is right, and 
I published my change of opinion in a paper on Some recent Changes 
in the Nomenclature of West Lidian corals.* Therefore, I now 
believe that Madrepora acropora Linnaeus should be considered as 
imdeterminable and that the name should be dropped from coral 
nomenclature. 

The type-specimen of Madrepora faveolata EUis and Solander is 
preserved in the Hunterian Museum of the University of Glasgow, 
where I have seen it, and Prof. Graham Kerr has kindly sent me a 
photograph. It is a worn specimen, considerably infiltrated with 
calcium carbonate, and is probably the same as OrhiceUa annularis, 

Astraea {OrhiceUa) steUulata' Dana has been carefidly redescribed by 
Professor Verrill from Dana^s types, which are preserved in the Yale 
University Museum. The following is his description: 

They ' are beach-worn specimens of a true OrhiceUa, mcro or less infiltrated with 
calcium carbonate, to which the unusual solidity of the walls and exothca, in some 

» iCeol. Soc. London Quart. Journ., vol. 51, p. 272, 1895. 

« Hist. nat. CoraD., vol. 2, p. 477 1857. 

t Conn. Acad. Arts and Sci. Trans., vol. 11, p. 94, 1902. 

* Bio!. Soc. Washington Proc., vol. 15, p. 56, 1902. 

• Conn. Acad. Arts and Sc-i. Trans., vol. 11, p. 96, pi. 15 ,fig. 2, 1902. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 373 

pckrts, as seen in sections figured by Dana, seem to be partly due. In other parts the 
»t:aructure is nearly as in 0. annUlariSf to which it probably belongs, though there are 
[Uiferences in the sections not due to infiltration. Its septal arrangement Is the same 
aiS in ordinary specimens of the latter, those of the third cycle being distinct, but 
Ei^urrow and thin. The borders of the calicles seem to have been but little raised, and 
t.li.e septa rather thinner than usual, and not much exsert, but the poor condition of 
tkxe specimens renders these characters rather uncertain. 

The calicles are rather smaller (2 to 2.5 nmi. in diameter) than is usual in 0. annu- 
laris. The thin septa are in three regular cycles; those of the third cycle are very 
t^liin and reach only one-fourth or one-third to the columella, which is well developed. 
Tlie septa are a little thickened at the wall; their faces are only slightly granulated. 
ITliere are a few, irr^:ular, small teeth on their inner edges where best preserved; 
xipper ends are all worn off; some have a paliform tooth at the base. The costae are 
-well developed, inosculating, with irregular exothecal dissepiments between them, 
£t8 in O. annularis. But in some vertical sections the walls appear as narrow, solid 
structures (where unaltered); in the sections the columella region is loosely filled 
Twith stout ascending trabeculae; the endotheca consists of small, very thin, nearly 
horizontal dissepiments, inclining downward a little, and often in two series. No. 
4266. 

Their origin is imcertain, but it appears to be West Indian. They are in the same 
beach-wom state as several other types of West Indian corals studied by Professor 
Dana. Apparently most West Indian corals, in good condition, were scarce in 
American museums at the time when he wrote his great work. 

It appears to be a small or somewhat dwarfed variety of 0. annularis. I have seen 
freeh specimens of a similar variety from the Florida reefs. 

This may well be identical with M. stellulata Ellis and Solander, but the latter 
can not be determined with any certainty from the figure, which represents a badly 
worn specimen. Its calicles, as figured, are mostly even smaller than in Dana's type, 
and somewhat unequal in size; the walls appear to be as solid as in the latter; the 
calicles project slightly as in annularis; 12 to 15 septa are figured, all perfect; colu- 
mella is as in annukaris. There is much more reason for calling this a variety of 0, 
annularis than there is for identifying it with Solenastraea hyades, as Gregory has done. 
There is no evidence that it is a Solenastraea, 

Fortunately Dana's OrhiceUa sUUulata is a synonym of 0. anmi- 
laris and is not even of varietal importance. Professor Verrill says, 
''This may well be identical with M, steUulata Ellis and Solander," 
an opinion from which I emphatically dissent. The figures of EUis 
and Solander are of a SoUnastrea (Nat. Hist. Zooph., pi. 63, figs. 3, 4); 
the costae do not continue from one calice to those of adjacent cahces, 
and the exotheca, as is shown by the side of figure 3, is typical of 
Solenastrea. Furthermore, in the description of the species it is 
stated, ''interstitiis planiuscuhs scabriusculis,'' the intercoraUite 
areas are not "radiate^* as in anrmlaris. The Heliastraea steTMlata 
of Milne Edwards and Haime (see pi. 80, figs. 7, 7a, 76) is not the 
Madrepora steUulata of ElUs and Solander; it is probably the same 
as OrbiceOa annularis. 

There is much doubt about the CypTiastraea oblUa Duchassaing and 
Michelotti. The following is the original description: 

Esp^e arrondie, avec des ^toiles arrondiee et k bord iin pen eUy6: c6tes rares, 
I^esque confluentes; les intervaUes de Tune k I'autre 6toile sent gamies de granu- 
lations; la columelle est grande et papilleuse. 



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374 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

La Cyphastrea oblUa a lee bords moinfi el4v^, et lee cIouods plus debordantes qce 
celles de la Cyph. microphthaXma qui sont aussi gamies d'une petite dent subptli- 
forme qui manque dans la Cyph. oblita. St. Thomas. 

I found in the Museum of NatuiiEtl History at Turin a specimen 
labeled '' Cyphastrea oblUa,^* It is a specimen of OrHeena annuiarit. 
Another specimen bearing the same label, seen in the Museum d'Eb- 
toire Naturelle at Paris, is a Solenastrea.^ The latter is a rounded 
head with a greater diameter of about 130 mm. The calices range 
in diameter from 2 to 3 mm.; distance apart from somewhat less to 
sUghtlj more than 1 mm., occasionally 2 mm. Margins of the calices 
marked by a slightly raised rim. Costae insignificant, occasionally 
extending from one calice to the next. Septa in three complete 
cycles, primaries and secondaries reaching the columella; tertiaries 
shorter, with inner edges free, i. e., not fused to the sides of a lower 
cycle. PaU variable in development; in some calices they are lai^. 
flattened above, before all septa except the last cycle; in others, 
several teeth indicate the position of a palus. Columella, lax and 
papillary. This specimen is the same as the Helia^ruea ahdiia 
Duchassaing and Michelotti. 

The original description of Gyphastrea oblita is not adequate for 
identification. One of the specimens from the Duchassaing and 
Michelotti collection is OrhiceUa annvlaris, the other the same is their 
own Heliastraea dbdUa. Because the Paris specimen is probably the 
type I am placing the species in the synonymy of Sclenastrea hour- 
noni M. Edwards and Haime (see p. 400). 

Heliastraea rotvlosa Duchassing and Michelotti is a growth form of 
0. annularis, judging by the description. I did not find the type in 
Turin. 

The specimens determined by Duchassing and Michelotti as 
Hdiasiraea acropora (Linnaeus) and H. lamarcki Milne Eklwards and 
Haime are, according to specimens bearing those names in the 
Museum of Natural History at Turin, referable to OrhiceUa annular\$. 

The type of Duncan's CypTuistraea cosiata from Barbuda is pre- 
served in the (jreological Society of London, and I studied it there. 
The specimen shows no noteworthy variation from the usual OrhieeJh 
annularis, except that its calices are from 3 to 4 mm. in diameter, 
usually 3.6 mm. Another specimen, from Santo Domingo, labeled 
CypTiasiraea cosiata is a Solenastrea. The specimens determined by 
Gregory as C. cosiata were studied in the British Museun of Natural 
History; they are 0. annularis, 

Astraea harbadensis Duncan is a specimen of 0, annularis from the 
Pleistocene reefs of Barbados. 



* lUustratioDS of this specimen have been published by me in U. S. Geol. Sarv. Prof. Pap. W-T, pi. 9B. 
ftRS. 3, 3a, 1917. 



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GEOLOGY AND PALEONTOLOGY OP THE CANAL ZONE* 375 

Gregory refers Hdiastraea dltissima Duncan to the synonymy of 
this species, but I doubt the correctness of his conclusion and am 
treating it as valid. 

Oeologic distribtUion. — Pleistocene and Recent, throughout the 
elevated reef areas of the West Indies, eastern Central America, and 
Florida. 

Duncan * has listed Astrctea barbadensisy one of the synonyms of 
O- annularis, from the "marl formation" of Antigua, remarking that it 
\3 ''greatly altered by f ossihzation ; the calicular surface is subplane, 
and the cahces are seen as prominent columnar casts." Should 
Dimcim's identification be correct, the geologic range of 0. annviaria 
extends from Oligocene time to the present. Mr. R. T. Hill obtained 
in Antigua a silicified specimen that looks like 0, annvlaris, but I am 
not sure that it is that species. 

Costa Rica, station 4269, Port Limon, collected by Doctor Wailes 
in beds referred to the PUocene. There are three dissociated coral- 
lites which have the general characters of OrbiceUa annularis, but are 
not absolutely typical, for the primary septa are appreciably but not 
strikingly thicker than the secondaries. They are, therefore, somewhat 
intermediate between typical examples of the species and OrbiceUa 
limbata (Duncan). 

2, ORBICELLA LIMBATA (Dwoui). 

Plate 85, figs. 1, la, ^, 2a, 25, 3, 4, 4a. 

1863. Pkyllocoenia aculpta var. iegula Duncan, Geol. Soc. London Quart. Joum., 
vol. 19, p. 432. 

1863. Phyllocoenia limbata Duncan, Geol. Soc. London Quart. Joum., vol. 19, 

p. 433. 

1864. Plesiastraea ramea Duncan, Geol. Soc. London Quart. Joum., vol. 20, p. 

39, pi. 5, figs, la, 16. 
1866. Pkyllocoenia limbata Duchassaino and Michelotti, Sup. M4m. Corall. 

Antilles, p. 76 (of reprint). 
1866. Plenastraea ramea Duchassaino and Michblotti, Sup. M6m. Corall. 

AntiUes, p. 87 (of reprint). 
1868. Phyllocoenia limbata Duncan, Geol. Soc. London Quart. Joum., vol. 24, 

p. 23. 
1868. Plesiastraea ramea Duncan, Geol. Soc. London Quart. Joum., vol. 24, p. 25. 
1870. Phyllocoenia limbata Duchassaino, Rev. Zooph. et Spong. Antilles, p. 28. 
1870. Plesiastraea rtrniea Duchassaino, Rev. Zooph. et. Spong. Antilles, p. 30. 

Original description of Phyllocoenia limbcUa:^ 

Corallum in the shape of Stylina limbata Edwards and Haime. Stem large 
and cylindrical. CoralUtes numerous, irregularly placed. Calicee separated by 
much coenenchyma, circular and but slightly elevated. Costae covwing much 
surface. Slightly dentate where they approach, and turning aside from those of oth«r 
calices; they are not c<Hitinuous, not very prominent, and slightly granular. Septa 
not projecting far inwards, laminae granular; their upper margin is neither incised nor 
dentate; in six systems of generally three cycles, though occasionally of four. Pri- 

1 Oeol. Soo. London Quart. Joom., vol. 19, 1863, p. 433. 
37149— 19— Bull. 103 13 



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376 BULLBTIK 103, UNITED STATES NATIONAL MUSEUM. 

mary septa largest. Columella rudimentary. Endotheea abundant. Diameter oi 
calice, with coetae, one-fifth inch [5 mm.]. 

The deficient columella is the only point in which this species differs from yfadrt- 
pora limkata Goldfuas, which has been determined by Milne Edwards to he a Stylira 

From the yellow shale of San Domingo. Coll. Geol. Soc. 

Original description of Plesiastraea ramea: ^ 

Corallum in gibbous mnosoo or more or less cylindrical processes with irregulir 
swellings. Calices distant, very slightly exsert, circular, and unequal in aize 
Septa thick at the wall, thin internally, unequal in size, according to the 
orders; finely dentate above, but sparely granular laterally. In six sN^stems of three 
cycles, with occasionally an additional order in one- half of a system. P^ very 
small. Coliunella lax, papillated, and small. Fossa mediately deep. Costae veil 
developed, subequal, and marked by three or four dentate projections; they are evi- 
dently covered with a fine epitheca, which is not granular; where the epitheca is wofB 
the costae are seen to be smaller, the tertiary being much smaller than the otheo: &U 
project, however. Exotheca moderately developed and often becoming indurated. 
Endothccal dissepiments fragile, but horizontal and frequent. Height, some inches; 
diameter of branehes 1 inch, more or less; diameter of coralUtes four-thirtieths inch 
[3.3 mm.]; distance between corallites about one-tenth inch [2.5 mm.]. 

From the silt of the Sandstone plain, San Domingo. Coll. Geol. Soc. 

I examined Duncan's types of PhyUocoenia limbaia and Plesiasiraec 
ramea in the Geological Society of London and made a note that the 
latter, except that its septa are broken down and the calices have a 
hollowed-out appearance, is the same as the former. 

In my Some fossil Corals from the elevated Reefs of Curasao. 
Arube, and Bonaire, and my Stony Corals of the Porto Kican 
waters, I placed in the synonymy of OrhiceUa acropora (=0. 
annularis), the three names of Dimcan, cited above, considering the 
specimens to which they were applied as growth forms of that species. 
More detailed studies, subsequently made, have led me to beliere 
that I was mistaken in that course. This coral is very similar to 0, 
annularis. However, there appear to be two constant differwices— 
namely, the primary septa within the calices are imiformly thicker 
and usually longer than the secondaries (this lesser development of 
the secondaries is not occasional as in O. annularis but constant) and 
small, but distinctly developed, pali occur before the primary and 
secondary septa. 

I have for study one specimen from Duncan's original material, 
labeled Plesiastraea ramea Duncan, No. 155273, U.S.N.M., kindly 
sent to the United States National Museum by the authorities of the 
Geological Society of London (see pi. 85, figs. 1, la); 10 specimens 
belonging to the Museum of Comparative Zoology, 4 specimens 
collected by Miss C. J. Maury in Santo Domingo, and material 
obtained by myself near Santiago, Cuba. The first specimen is not 
in very good condition for study, and does not fit Duncan's descrip- 
tion well. The Museum of Comparative Zoology specimens, however, 

>060l. Soc. London. Quart. Jour., vol. 10, 1863, p. 421. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 377 

fit exactly, omitting the remarks about the costae being covered by 
epitheca. The figures presented on plate 85, figs. 2, 2a, 2h, and 3, 
are based on these specimens. 

PhyUocoenia sculpta var. (egvla Duncan.* As I do not find Dun- 
o€Ui*s description of this coral satisfactory, and as the authorities of 
tlie Geological Society of London -have kindly sent one of the original 
specimens to the United States National Museum (No. 155274), 
(see pi. 86, figs. 4, 4a), I submit the following description: 

Corallum, a rather thick foUimi; the specimen here described is 
unfortunately broken on all its edges, its original dimensions are there- 
fore imknown. Its present length is 62 mm. ; width 40 mm. ; greatest 
thickness, 15.5 mm.; thickness near outer edge 5.5 mm. Base in- 
vested with a coarsely wrinkled epitheca. 

The caUcular margins are on the same level as the flat exothecal 
surfaces, or are veiy slightly raised. In form the calices are circular 
or somewhat deformed. Diameter from about 2 mm., to 2.5 by 
*3.25 mm.; distance apart, from 1 to 3 mm. IntercoraUite areas with 
costae, beaded on the edges, equal or alternating in size, correspond- 
ing to all septa, those of one calice meeting those of the adjoining 
calices. 

Septa usually in three complete cycles, primaries and secondaries 
li rger, and usually thicker, than the tertiaries, primaries average 
la/ger than the secondaries. All the primaries and most of the second- 
ariv\s reach the columella. 

Columella trabecular. 

Locality and geologic occurrence. — Nivaj^ shale, Santo Domingo, t. 
Duncan. 

Miss Maury obtained specimens in Santo Domingo as follows: 

Rio Cana, zone H, associated with Pla^cocyathus, new species, 
Siylophora granuUUa Duncan, AntiUia hilohata Dimcan, OrhiceHa lain- 
bridgensis Vaughan?, Solenastrea boumoni M. Edwards and Haime, 
SyzygophyUia gregorii (Vaughan), and Siderasirea siderea (Ellis and 
Solander). Rio Gurabo, zone D, associated with StylopJiora affinis 
Dimcan, Madrads decadis (Lyman), PociUopora cra^soramoaa Dim- 
can, StepJumocoenia intersepta (Esper), OrbiceUa cavernosa var. 
cylindrica (Duncan), SyzygophyUia dentata (Dimcan) ; zone not stated, 
associated with PodUopora crassoramosa, TJiysanus grandis (Dimcan), 
and SyzygophyUia dentata (Duncan). 

I collected in 1901 a fine specimen of this species east of La Cruz, 
at the crossing of the highway from Santiago to the Morro over the 
railroad, near Santiago, Cuba. Other corals collected there, including 
Siylophora species (probably S. affinis Duncan), Solenasirea hour- 
noni M. Edwards and Haime, a species of Thysanus (aff. T. exeen- 
iricuA Duncan), Siderasirea siderea (EUis and Solander), and Ooniopora 

> Geol. Soc. London Quart. Jouro., vol. 19, p. 432,a8d3. 



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878 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

jctcobiana Vaughan, indicate similarity in horison with zone D of the 
Rio Gurabo section. 



t. ORBICBLLA IMPBRATORIS. new i 

Plate 86, figs. 2, 3, 4, 5. 

Corallum forming rounded mass^ 16 cm. or more in diameter. 

Calices in the type-specimen are not much elevated but have a dis- 
tinct, somewhat raised wall; in other specimens the corallites may 
project as much as 2.5 to 3 mm. Calicular diameter, 3.5 to 5 mm.: 
distance between calices, from 2 to 3.5 mm. Corallites joined by 
prominent, rather thin, distant costae, which correspond either to 
all cycles of septa or to the primaries and secondaries. 

Septa, typically in three complete cycles; the 6 primaries promi- 
nent, thicker than the members of the higher cycles, and extend 
to the columella; the secondaries usually do not reach the columella, 
only about half as long as the primaries; tertiaries shorter and thin- 
ner than the secondaries. The septa are usually distant in the wall. 
The third cycle of septa is incomplete in some calices; while in large 
calices a few secondaries may reach oi almost reach the colimieUa. 

Columella formed by the fusion of the inner edges of the primary 
septa. 

Endotheca well developed as dissepiments. Exotheca well devel- 
oped between the strong costae, about 3 dissepiments within 1.5 mm. 

LocoMHes and geologic occurrence. — Canal 2iOne, Panama, in the 
Emperador limestone, at stations 6015 and 6016, quarries in Empire, 
and 6017, one mile from Empire toward Las Cascadas, collected by 
T. W. Vaughan and D. F. MacDonald; station 6256, in the Emperador 
limestone, 1} miles south of Miraflores, collected by D. F. MacDonald. 

Cuba, station 3450, 4 miles north of the City of Pinar del 
Rio, and station 3451, one-half mile west of Cienaga railroad station, 
near Habana, collected by T. W. Vaughan; station 3566, Bejucal, 
collected by Arthur C. Spencer; station 7544, Rio Yateras, near 
Guantanamo, collected by O. E. Meinzer. N. H: Darton, collected 
at station 7664, on the north slope of La Piedra, northeast of Jamaica, 
which is northeast of Guantanamo, a specimen of QrbiceUa apparently 
referable to this species. The calices are rather large, 7 nmi. in 
diameter, and nearly all of the secondary septa reach the colimiella. 
It seems very near O. anthUarum. The specimens from Cienaga, 
Cuba (pi. 86, fig. 5), is illustrated as well as the cotypes from Panama. 

Anguilla, stations 6893 and 6967, Crocus Bay, collected by 
T. W. Vaughan. 

Cotypes,— iiio, 324884, 324902, 324903, U. S. N. M. 

Paratype.—^o, 324878, U. S. N. M. 

This species is distinguished by the small size of its calices, its 
prominent costae, its 6 long septa, with intermediate septa shorter 
according to cycle. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 379 

4. ORBICELLA ANTHXABUM (Dwicui). 

1863. Astrcbea antUlarum Duncan, GeoL So<^. London Quart. Joum., voL 19, p. 
443. 

1866. Astraea antUlarum Duchassaing and Miciielotti, Sup. CoralL Antilles, 

p. 86 (of reprint). 

1867. Heliasiraea antUlarum Duncan, Geol. Soc. London Quart. Joiirn., vol. 24, 

p. 24. 
1870. Heliastraea antUlarum Duchassainq, Rev. Zooph. Antilles, p. 30. 

Thi8 coral was doubtfully referred by me * to the synonymy of 
OrbiceUa cavernosa; but, as there is doubt, it is here accorded specific 
treatment. 

Original description, — "A specimen in the form of a rolled flint, 
found with silicified wood, has the corallum large, tall, probably 
resmbling in shape that of the San Domingan A. exothecata. Coral- 
lites close, unequal in size, but quite distinct; the transverse section 
shows them to be circular in outline. Septa in six systems of three 
cycles. The primary and secondary septa are nearly equal, and reach 
to the columella; the tertiary are small and straight; all are slender, 
wide apart, and very distinct. Costae tolerably developed, subequal. 
Walls moderately developed, by no means strong. Columella small, 
and occupying a small space. Endotheca greatly developed, vesicu- 
lar, and forming cells between the septa. Exotheca well developed; 
large cells broad, others squarer, with shelving dissepiments. Diam- 
eter of the coraUites three-tenths inch [7.5 mm.]. The interspaces are 
filled with opalescent or porcellanous silica; sclerenchyma often 
destroyed. Coll. Geol. Soc.*' 

Locality. — Montserrat. 

Duncan ' lists a coral as Astrasa antiUarum variety, and makes 
the following note: **With more distant calices than the type, pro- 
duced costae, and a less perfect development of the third septal 
cycle. The exact locality is not known, but the coral, from its 
mineralogical characters, appears to have been obtained from An- 
tigua. Brit. Mus. " 

Regarding the apparent absence of a fourth cycle of septa, it will 
be noted that as they are often very small and may be broken away, 
something especially likely to occiu: in worn specimens such as 
fossils usually are, they may have been present, but were subse- 
quently destroyed. The size ol the calices, 7.5 mm. in diameter, 
suggests the presence of quaternary septa. Additional material 
from Montserrat is needed to solve the question. 

5. ORUCELLA ALTlSSUtf A (Dwmui). 

1867. Heliastraea altissima Duncan, Geol. Soc. London Quart. Joiurn., vol. 24, 
pp. 12, 24, pi. 2, fig. 3. 

Original description, — " The corallum is very massive and tall, and 
its upper surface is subplane and wider than the base. The calices 

1 Qeologisch. Relohs. ICos. Leiden SammL, ser. 2, vol. 2, pt. 1, p. 28, 1901. 
* Geol. Soo. London Quart. Jonrn., vol. 20, p. 30, pi. 4, fig. 2. 



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380 BULLETIN ia3, UNITED STATES NATIONAL MUSEUM. 

are barely above the common surface, they are circular, but occa- 
sionally deformed, and they are slightly imequal in size. The calicular 
fossa is shallow, and the calicular margins are broader than the 
septa. The columella is small, distinct, lax, and parietal. The 
costae are well marked, unequal, and rarely touch, and they are 
thicker than the septa. The costae of the highest order are well 
developed, and contrast with their rudimentary septa. The septa 
are deUcate, they arc thinner midway than elsewhere, and those 
which reach the columella have a palif orm tooth ; they are not exsert, 
and are only slightly dentate. The septa are very irregular in their 
arrangement. There are six systems, and in most of them there 
are three cycles with or without a part of a fourth in one-half of the 
system, so that there are constantly six septa in a system instead 
of eight. The endotheca is well developed; and the dissepiments 
arc close, stout, and nearly horizontally parallel. The exotheca is 
abundant, forming small cells with arched outlines. Height of 
corallum 6-8 inches. Diameter of caUces two-tenths inch [ = 5 mm.]." 

Locality. —?>t, Croix, Trinidad. 

Gregory ^ places Duncan's Heliastraea altissima in the synonymy 
of Orbicella acropora (Linnaeus), without giving his reason. He may 
be right, but the caUces are large for 0, acropora (here called 0. 
aniivlaris), and judging from the presence of quaternary septa it is 
almost certainly distinct. According to Duncan's figure every other 
septum reaches the coliunella, a septal arrangement which is one of 
the characteristics of 0. annularis. I did not see the type in London, 
and think that until it is restudied or additional material has been 
collected at the type locality, it will not be possible to reach a positive 
decision as to the validity of the species. 

t. ORBICELLA CAVERNOSA (lisMeM). 
Plate 87/figB. 1, la, lb, Ic; plate 88, ige. 1, 2, 3, 3c, 36. 

1766. Madrepora cavernosa Linnaeus, Syet. Nat., ed. 12, vol. 1, p. 1276. 

1901. Orbicella cavernosa Vauohan, Geolog. Reichs. Mub. Leiden Samml., ser. 

2, vol. 2, p. 27 (Synonymy with exceptions noted below). 

1902. Orbicella cavernosa Verrill, Conn. Acad. Arte and Sci. Trans., vol. 12, p. 

102. 

1915. Orbicella cavernosa Vauohan, Washington Acad. Sci. Journ., vol. 5, p. 596. 

1916. Orbicella cavernosa Vauohan, Carafe Inst. Washington Year Book No. 14, 

p. 227. 

In my first paper referred to in the synonymy I placed Asiraea en- 
dothecata, Astraea cylindrica, and Astraea hrevis of Dimcan in the 
synonymy of this species. A, endoihecata and A. cylindrica now 
seem to me to deserve varietal recognition and A. brevis should be 
treated as a valid species imtil additional information concerning it 
is available. Duncan's Astraea aniiguensis, which I doubtfully 

I Oeol. Soc. Land<m Quart. Joatn., voL 51, p. 272, 1896. 

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GEOLOGY AND PALEONTOLOGY OP THE OANAL ZONE. 381 

placed in the synonymy of 0. cavernosa j has the general appearance 
of OrhiceUa, but it is a fungid coral and is referred to the genus 
C*yathomorpha. Astraea antiUarum, given by me as doubtfully a 
synonym of 0. cavernosa^ should for the present at least be treated 
3^ a vaUd species. 

As so many of the species related to 0. caA)emo8a must be con- 
sidered in this paper, it is desirable to describe all those members of 
trte group found in the American Tertiary formations and now 
living in the western Atlantic Ocean. The systematic rank of the 
forms described on the following pages is open to debate, and I wish 
liere to express my recognition of other methods of treatment. As 
the forms, whether they bo designated '^species/* '* subspecies, '' 
* ' variations, ' ' or merely ' ^ variants, ' ' exist, and as they have geographic 
and geologic significance, they should be discriminated and char- 
acterized. In comparison with these desiderata nomenclatorial 
considerations are of secondary importance. 

OrbiceUa cavernosa is so variable that great difficulty has been 
experienced in constructing an intelligible description. A very 
interesting specimen, obtained by Prof. J. E. Duerden in Jamaica 
and presented by him to the United States National Museum, will 
first be described in detail, as it shows within itself a wide range of 
variation and indicates the lines of variation of other specimens 
more constant in their character (see pL 87, figs. 1, la, lb, Ic). 

The coralliun is oblong; upper surface convex but not xmiformly 
arched or domed; base epithecate. Length, 25 cm.; breadth, 20 
cm.; thickness, 11.3 cm. 

The specimen has two different kinds of calices. Those of one 
kind are rather distant, protuberant, and have subequal, not very 
tall, thick, dentate costae (pi. 87, fig. 1). The transverse outline is 
circular or broadly eUiptical, diameter between thecal simmiits 
8 mm. ; one of the elliptical calices has a greater diameter of 11 mm., 
lesser about 9 mm. The costae are about 1 mm. tall. The distance 
apart, measured between the outer costal edges, is from almost 
contiguous to 6 mm. The free limb of the corallite is subcylindrical 
and projects between 6 and 7 mm. The caUces, as is shown by 
plate 87, figures 1, la, are not imiformly distributed, and vary m 
size, form, and prominence. 

In a fully developed calice there are 48 septa, every other one 
extending to and fusing to the columella. All the septa, particularly 
the principals, are rather thick. The margins are d^itate, within 
the calicular cavity, they fall abruptly to the bottoms of the caUces, 
which are 3 to 4 mm. deep, and there the principals extend to the 
columella. There are septal teeth around the periphery of the 
columella but they are not in the form of well-developed pali or 
paliform lobes. 



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382 BULLETIN 103, UNITED STATES NATIONAL MUSEUJC. 

The columella is large, trabecular, with a papillate upper surface; 
diameter, as much as 4 mm. The columellar elements axe rather 
often twisted and present a whorled appearance. 

Endo thecal dissepiments rather deUcate; exotheca, coarse and 
very vesicular. 

The calices of the other kind (pi. 87, fig. la) in their typical devel- 
opment are smaller than those above described, their edges are 
only slightly elevated, and the septa and costae are decidedly thin 
and exsert. Diameter of the calices from 5 to 8 mm. ; septal margins 
2 mm. tall. 

The differences between these two kinds of calico are so gre^t 
that it seems scarcely possible that they could belong to the same 
species; however, they occur on the same corallum where j>erfect 
intergradation can be traced. 

Pourtalfts, as far back as 1871,* published the following important 
notes on this species: 

There is considerable variation among the specimens from Florida in the Museum of 
Comparative Zoology, enough apparently to warrant placing them among the three 
species mentioned in the synonymy; but by car^ully examining the different part? 
of each specimen, passages from one to the other can be found. Thus young polypi- 
doms, expanding rapidly laterally and with rather distant polyps, appear at first to 
differ considerably from strongly convex ones with crowded OSdices; the ooetae are 
larger, flatter, and less sharply denticulate, and the border of the calicles leas ele\'ated. 

The size of the calicles, relied on to divide the genus into groups by Milne Edwards 
and Haime, is a very uncertain character; one specimen has in one part the calicles 
varying from 3.5 to 4 mm., in another from 7 to 8 mm. The same specimen has in 
some parts the contiguous walls united solidly, with very few or no exothecal cells, in 
others separated by an abundant cellular exotheca. In worn specimens the last cycle 
disappears first, for that reason probably Orbicella ( Madrepora) radiata Ellis has been 
characterized by the Milne Edwards and Haime as having but three cycles. 

Venill gives the following description:' 

Much of the confusion in regard to the name of this species is due to the fact that it 
was generally described and figured from badly beach-worn specimens by the eariier 
writers. Such specimens havte the septa and calicles worn away and the hard ex- 
otheca thus becomes prominent around the excavate calicles, so as to greatly c^iange 
the appearance of the coral. Another cause is the rather wide variations in the aixe 
of the calicles. 

The normal or average specimens have the calicles about 6 to 8 nun. in dianieter. 
but occasionally a specimen occurs in which part or all of them may be 9-10 mm., or 
rarely, even 11 mm. in diameter. Sometimes, on crowded parts of large specimens. 
the diameter may be only 4 to 5 mm. The degree of elevation of the calicles is also 
more or less variable on a single specimen. 

The calicles may be pretty close together, where crowded, but in other cases they 
are separated by spaces of 4 to 6 mm. or more. The costae are usually well developed 
as denticulated, rounded, radial ribs, usually 48 in number. 

The septa are generally about 48, arranged in four regular cycles, but several of 
those of the last cycle are often rudimentary or lacking, reducing the niunb^ to 40-44 

1 Mus. Comp. Zool. 111. Cat., No. 4, p. 70. 

* Trans. Cono. Acad. Arts and Sci., vol. U, pp. 102, 103, 1902. 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 383 

T'lx^iy differ in breadth and thickness according to the cycles; those of the last cycle 
aje "very thin and often bend toward and join those of the third cycle. The principal 
Bep^s are exsert, denticulated, and thickened at the wall. The columella is usually 
xvell developed and broad. The paliform teeth are distinct? but not very prdtaiinent. 
X t, sometimes forms hemispherical masses 4 to 5 feet or more in diamet^. 

'Plus species appears to be rare at the Bermudas, and probably occurs only on the 
oxxtermost reefs. The only specimen seen by me from there was from near the North 
Rooks. (Centennial collection.) It is a hemisphere about 11 inches in diameter, of 
tbe typical form. It is common on the Florida reefs and throughout the West Indies, 
ri3liia, Brazil; (YaleMu8.);=var.^«rta,nov., with elevated corallites; roughly serrate* 
thin costae and septa; dalicles deep, 5-6 mm. broad; septa narrow, perpendicular 
i^lthin, usually 40-44. 

The description of the Jamaican specimen, when taken in connec- 
trion with thfe notes by Pourtalfes and Verrill, gives a goodidea of the 
ci^tent of the variation of the species except in one particular, that of 
tlie septal arrangement. The normal, f uUy developed cahces have four 
eomplete cycles of septa; however, sometimes the foiurth cycle may 
not be complete while at others there may be a few qtdnaries. In 
tlie recent specimens the tertiaries usually, but not invariably, 
extend to the columella. 

The characters common to all of the specimens may be briefly 
summarized as follows: 

Corallum massive, base epithecate, upper surface flat, irregularly 
convex, or domed. Calices more or less elevated, diameter from 5 to 
11 mm., externally costate, oostae normally subequal. Septa nor- 
mally in four complete cycles, the members of the first three cycles 
extend to the columella, but the fourth may not be complete, and 
sometimes there may be a few quinaries. Columellar trabecular, 
well developed, large, with a papillary upper surface. 

Remarks on the syrionomy of 0. ca/vemosa. — The names 0. radiata 
CEIUis and Solander), 0. argus (Lamarck), 0. conferta (Milne Edwards 
and Haime), and 0. cavernosa var. Mrta Verrill, are definitely placed 
in the synonymy of 0, cavernosa, and it is thought probable that 
O. brazUiana Verrill, should be referred Jbo it. These names will be 
discussed seriatim. 

Gregory appUes 0. radiata to this species, as he considers the lin- 
naean defmition of Madr&pora caA)emosa insuflScient, an opinion with 
which I do not agree. All the linnaean descriptions are unsatisfactory, 
but in this instance Linnaeus refers to the figures of Seba, he places the 
Madrepora astroites of Pallas in its synonymy, and he states ** Habitat 
in O. Americano." Taking all things together, the original diagnosis 
with the references seem to me sufficient for purposes of identifica- 
tion — in fact, the brief Latin description is not bad. 0. radiata was 
supposed to diifer from 0, cavernosa by possessing only three cycles 
of septa. Pourtalfis states, in the quotation already made from him, 
that ''In worn specimens the last cycle disappears first; for that 
reason probably Orbicdla (Madrepora) radiata Ellis and Solander has 



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384 BULLETIN 103, UNITED STATES NATIONAL MU8EX7H. 

been characterized by Milne Edwards and Haime as having but 
three cycles;" 

Lamarck's Astrea argus is a new name for the Madrepora cavernosa 
Elsper. The reason for his giving it is not evident. 

The specimen identified by Ehrenberg as Ezplanaria argus, which 
is the type of Milne Edwards and Haime Astrea conferta, is in the 
Berlin Museum fiir Naturkunde, and the following notes are based 
upon it: 

The specimen is much worn and is apparently somewhat fossilized. 
The caUces are not regularly rounded, but frequently are of irregular 
polygonal outUne. The greater diameter of an average cahce is 
8.5 mm.; lesser 7 mm. Thickness of wall between the ealices is 2.5 
mm. In one caUce there were 21 large and 21 smaller septa; there 
may be four complete cycles in some caUces. The columella is very 
large and vesicular and occupies the greater part of the coraUite 
cavity. Dissepiments abundant, about 13 to 5 mm.; they slope 
downward and inward. From reading the Pom*tal6s description 
quoted above, it will be evident that this is only a variety of 0. 
ca/vemo8a with crowded caUces. The ExpUmaria radiaia of Elhrenberg 
is the ordinary Hdiastraea cavernosa as figured by Milne Eklwards 
and Haime, except that the fourth cycle of septa may not always be 
complete. 

OrbiceUa davemoea var. compacta Vaughan (pi. 88, figs. 3, 3a, 36) 
has solid exotheca, low mammillate corallites, and equal costae. 
Recent on the Brazihan coast; lat. 12'' 48' S., long. 38*" W.; 27 
fathoms. 

Localities and geologic occurrence, — On the living and Pleistocene 
reefs of Florida, the West Indies, and the Caribbean side of Central 
America. There are beach worn or Pleistocene specimens from the 
Isthmus of Darien in the United States National Museum, collected 
by Dr. Van Patton. 

«A. ORBICELLA CAVEBNOSA rar. BNDOTHBCATA dhncaa). 
Plate 89, figs. 1, la. 

1863. Astraea endothecata Duncan, Geol. Soc. Lond. Quart. Journ., vol. 19, 

p. 434, pi. 15, figs. 7a, 76. 
1868. Heliastraea endothecata Duncan, Geol. Soc. Ix)ndon Quart. Joum., vol. 24, 

p. 24. 

The corallite walls are thick; costae strongly alternating in size; 
the last cycle are small and thin, and there appear to be no septa 
corresponding to them; occasionally there is a rudimentary septum 
of the fourth cycle. The last cycle of septa may have been broken 
oflE; or the wall, because of subsequent thickening, may have included 
their inner ends; all other septa, with rare exceptions, extend to the 
large, woD developed columella. Diamet-er of coraUites from 8 to 



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GEOLOGY AND PALEONTOLOGY OF THE CANAL ZONE. 385 

.0 mm. Type in the Geological Society of London; duplicate in the 
Jriited States National Museum (No. 155276). The preceding re- 
xia,x*ks are based on the latter. 

LoccHides and geologic occurrence. — Type said to come from the 
NivajS shale of Santo Domingo. 

Costa Rica, station 4269, Port Limon; collected by Doctor Wailes 
in a bed of reputed Pliocene age. The size of the calices, and the 
costae, wall, and columella of the Port Limon specimen are as in var. 
endotJieccUa; but usually every other septum meets the columella; a 
cycle of small septa between the larger is clearly present. As it is 
probable that the last cycle of septa has been destroyed in the type of 
var. endotkecdtaf the presence of small septa between the larger would 
not indicate specific difference. The strongly developed costae with 
small ones between them are the same in both the type and the Port 
Limon specimens. 

The stratigraphic range of this variety, therefore, is from the 
XivajS shale (lower Miocene) to probably Pliocene. 

•b. ORBICELLA CAVERNOSA var. CYUNDRICA (Duncan). 

Plate 89, fig. 2. 

1863. Astraea cylindnca Duncan, Geol. Soc. London Quart. Journ., vol. 19, p. 

434, pi. 15, fig. 8. 
1868. Heliastraea cylindriea Duncan, Geol. Soc. London Quart. Journ., vol. 24, 

p. 24. 

This variety closely resembles var. endothscala. It has smaller 
corallites, 5 to 6 mm. in diameter; fewer septa, 12 to 16 principal 
septa, with from 1 to 3 smaller intermediate septa. Between each 
pair of larger s^pta on the mural smnmits around the calices is an 
intermediate rudimentary septum; the total number of septa is 
about 38. The costae corresponding to (he principal septa are 
strikingly prominent, while those corresponding to the rudimentary 
septa are very small or even obsolete. The calice is rather deep, 
about 2.5 mm. 

This coral may be only a growth facies of 0. endothecata. 

Localities and geologic occurrence. — Duncan type, in the Geological 
Society of London, comes from *'the tufaceous limestone^' of Santo 
Domingo; dupUcate specimen No. 155277, U.S.N.M. Miss C. J. 
Maiuy has recently collected the variety in Santo Domingo as fol- 
lows : 

Rio Gurabo, zone D, associated with Stylophora affinis Duncan, 
Madracis decactis (Lyman), PodUopora crassoramosa Duncan, 
StepJianocoenia intersepta (EJsper), OrhiceUa limhata (Duncan), Orbi- 
ceUa cavernosa (Linnaeus) var., SyzygophyUia dentata (Duncan), 
The single specimen collected is essentially typical, in fact it is a 
better specimen than Duncan's type. Cercado de Mao, without 



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386 BULLETIN 103, UNITED STATES NATIONAL MUSEUM. 

more specific information. The latter specimen has corallites wiih 
somewhat larger diameter, as much as 7.5, than those of typical 
specimens, diameter 4.5 to 5 mm., in that respect more doselj 
resembUng var. endothecatdy but there are no or only a few small 
costae between the large ones, and the septal characters are mort 
similar to those of var. cylindrica, 

Costa Rica "Colline en d6moUtion'^ Port Limon, No. 669, col- 
lection of H. Pittier. The specimens from Port Limon consist of 
two isolated corallites, which so closely resemble those of the type 
of var. cylindrica as not to need comment. Except in size, they art 
very similar to var. endotJiecata. 

7. ORBICELLA APERTA (VerrflU). 

Plate 89, fig. 3. 

1868. Helioitraea aperta Verrill, Conn. Acad. Arts and Sci. Trans., vol. 1, p. 356. 
1902. OrbiceUa aperta Verriix, Conn. Acad. Arts and Sci. Trans., vol. 2, p. Ift^, 
pi. 33, figs. 1, la. 

This species is especially characterized by having the principal 
septa, that is, those of the first, second, and third cycles, all of which 
oidinarily reach the columella, taller and thinner than is usual in 
0. c(wemo8a. At one time I was inclined to consider it only a varietr 
of OrbiceUa cavernosa, but comparisons of large suites of 0. cavernosa 
from Florida and the West Indies with a good suite of O. aperia 
from Brazil shows persistently recognizable differences. 

LocaiUies. — ^Abrolhos reefs, Bay of Bahia, and Island of Itaparica, 
Brazil. 

a. ORBICELLA BAINBIUI>GBNSIS» new spedM. 

Plate 90, figs. 1, la, 16, Ic. 

In growth form, general aspect of the corallum, and size of calices 
similar to OrbiceUa cavernosa. 

Calices 6 to 7 mm. in diameter; walls slope from caUcular margins 
to bottom of intercorallite areas; protuberant about 2 mm.; distance 
apart from 1.5 to 3.5 mm. 

Costae subequal, relatively thick, rather low, beaded on the 
edges, correspond to all septa, meet in the intercorallite depression. 

Septa in nearly four complete cycles, 10 to 12 septa, i. e., the 
primaries and most or all of the secondaries are thicker than the 
other septa, these and in some caUces a variable number of tertiaries 
extend to the columella. Usually the tertiary septa do not reach 
the coliunella, and the quaternaries are still shorter. The septa are 
distinctly of three sizes, even where the tertiaries reach the columella 
they are thinner than the members of the lower cycles. Septal 
margins dentate; distinct, rather wide, erect paliform lobes usual 
on the inner ends of the primaries and secondaries and in places 
on the tertiaries; on some septa instead of pahform lobes there are 



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GEOLOGY AKD PALBONTOLOGY OF THE OANAL ZONK. 387 

several teeldx with rounded upper ends. The youngest septa are 
Largely composed of ascending spines which are not completely fused . 
Columella relatively large, composed of septal trabeculae, upper 
surface coarsely papillate. 

Endothecal dissepiments highly developed, forming curved vesicles. 
Exotheca composed of successive, superposed but separated plat- 
forms extending between corallites (see pi. 90, fig. Ic). 

Localities and geologic occurrence. — Georgia, stations 3881, Blue. 
Spring, 4 miles below Bainbridge, and station 3883, Hales Landing, 
7 miles below Bainbridge, Flint River, Decatur County, collected by 
T. W. Vaughan; in the basal part of the Chattahoochee formation, just 
above the contact with the Ocala limestone. In the base of one 
sp>ecimen from station 3383 there is a cast of the surface of CerUhvam 
vaughani Dall, and there are several specimens of orbitoidal foram- 
inifera, one of which is clearly a species of Lepidocydina, Stations 
6085, Withlacoochee River, a few hundred yards below the Val- 
dosta Southern Railway bridge, and 6084, about 3 miles below the 
same bridge, Lowndes County, Georgia, collected by L. W. Stephen- 
son. 

Typ^.— No. 324881, U.S.N.M. 

Santo Domingan specimens, representing 'a very closely related if 
not identical species, were obtained by Miss C. J. Maury, on Rio 
Oana, in what she refers to as zone H, in association with a fauna 
representing the Bowden horizon, namely, Fhcocyatlma new species, 
Stylophora grawulata Duncan, Antittia bihbata Duncan, OrhiceUa Urn- 
bata (Duncan), SolenaMrea boumoni M. Edwards and Haime, Syzy- 
gophyUia gregorii (Vaughan), and SideraHrea siderea (Ellis and 
Solander). 

9. OBBICELLA COSTATA (Doncan). 

Plate 91, figs. 1, la, 2, 3, 3a; plate 92, figs. 1, 2, 3; plate 93, figs. 1, la. 

1863* Astraea costala Duncan, Geol. Soo. London Quart. Joum., vol. 19, p. 422, 

pi. 13, fig. 9. • 
1867. Heliastraea costala Duncan, Geol. Soc. Ix)ndon Quart. Joum., vol. 24, p. 24. 

Original description, — "The specimens of this species which I have 
examined present polished longitudinal and transverse sections of 
corallites, but I have seen no calices. CoralUtes long, parallel, some- 
times deformed, generally circular in transverse outline, not crowded, 
but close, varying in size. IntercoraUite spaces very distinct. Walls 
thin, not thicker than the delicate septa. Costae large alternately, 
both sizes equally produced; wedge-shaped at the wall, pointed, and 
often bent at the free end. Septa all deUcate and linear near the 
columella and in the middle; at the wall their base is narrower than 
that of the costae. They are arranged in six systems, the cycles 
being very irregular. In three systems there are three cycles, and 
in the rest an incomplete fourth; rarely there are two systems with 



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388 BULLETIN 103, UNITED BTATB8 NATIONAL MU8BUM. 

four complete cydee; the fourth and fifth orders often curve toward 
the third order. Lamellae rather cribriform, joining the cohimena 
by oblique processes. Columella lax, small, and formed by dissepi- 
ments from the septa and a central spongy mass. Endotheca Teiy 
abundant, vesicular, and horizontal, with four or five dissepiments 
in one-tenth inch [2.5 mm.]. Exotheca abundant, neariy equal to 
the endotheca. Reproduction by extra-caUcular budding. Diam- 
eter of the coraUites from three-tenths to seven-twentieths inch [7.5 
to 8.25 mm.]. 

'^This species is closely allied to the astraeans with great endo- 
thecal development, and especially to Astra^a vesiculosa Edwards 
and Haime, from Dax, as well as to A, antiUarum nob., and A. 
endothecaia nob.^' 

Locality.— ''The Marl of Antigua.'' 

Illustrations based on one of Duncan's original specimens, but not 
the type, are given on plate 91, figures 1, la. 

This species is represented in the Antigua formation of Antigua, 
the Pepino formation of Porto Rico, the Culebra formation of the 
Canal Zone, and in the marls and Umestone of Anguilla. The prin- 
cipal variation consists in the size of the calices. The minimum size 
of the calices in the Antiguan specimens (pi. 91, figs. 2, 3, 3a) is 
about 7 mm., which is about the average for the Porto Rican speci- 
mens (pi. 92, fig. 1), and the caUces of the specimens from the Canal 
Zone (pi. 92, fig. 2) average slightly smaller than those from Porto 
Rico, but the two sets of specimens differ very Uttle. In Anguilla 
(pi. 92, fig. 3;